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PROCEEDINGS

OF THE

California Academy of Sciences

Volume 43

SAN FRANCISCO

PUBLISHED BY THE ACADEMY

1982-1984

PUBLICATIONS COMMITTEE
Sheridan Warrick, Editor

Frank Almeda

Daphne G. Fautin

Tomio Iwamoto

Frank H. Talbot

(US ISSN 0068-547X)

The California Academy of Sciences

Golden Gate Park
San Francisco, California 94 1 1 8

PRINTED IN THE UNITED STATES OF AMERICA
BY ALLEN PRESS, INC., LAWRENCE, KANSAS

CONTENTS OF VOLUME 43

Pages

No. 1 . ROBERTS, TYSON R. The Southeast Asian freshwater pufferfish genus Chone-
rhinos (Tetraodontidae), with descriptions of new species. Published June 15,
1982 1-16

No. 2. McCosKER, JOHN E., AND JOHN E. RANDALL. Synonymies of Indian Ocean eels,
with the description of Gymnothorax enigmaticus, a moray previously known
as G. Ruppeli. Published June 15, 1982 17-24

No. 3. PULAWSKI, WOJCIECH J. New species of North American Tachysphex wasps

(Hymenoptera, Sphecidae). Published June 15, 1982 27-42

No. 4. ROBERTS, H. RADCLYFFE, AND CARLOS S. CARBONELL. A revision of the grass-
hopper genera Chromacris and Xestotrachelus(Orthoplera, Romaleidae, Roma-
leinae). Published November 4, 1982 43-58

No. 5. McCosKER, JOHN E. A new genus and two new species of remarkable Pacific
worm eels (Ophichthidae, subfamily Myrophinae). Published November 4,
1982 59-66

No. 6. RAINBOTH, WALTER J. Psilorhynchus gracilis, a new cyprinoid fish from the

Gangetic lowlands. Published July 6, 1983 67-76

No. 7. LE BOEUF, BURNEY J., DAVID AURICLES, RICHARD CONDIT, CLAUDIO Fox, ROBERT
GISINER, RIGOBERTO ROMERO, AND FRANCISCO SINSEL. Size and distribution of
the California sea lion population in Mexico. Published July 6, 1983 77-85

No. 8. TAYLOR, LEIGHTON R., L. J. V. COMPAGNO, AND PAUL J. STRUHSAKER.
Megamouth— a new species, genus, and family of lamnoid shark (Megachasma
pelagios, family Megachasmidae) from the Hawaiian Islands. Published July
6, 1983 87-110

No. 9. CRUMLY, CHARLES R. The cranial morphometry of Galapagos tortoises. Pub-
lished January 1 7, 1 984 111-121

No. 10. PULAWSKI, WOJCIECH J. The status of Trypoxylon figulus (Linnaeus, 1758),
medium De Beaumont, 1945, and minus De Beaumont, 1945 (Hymenoptera:
Sphecidae). Published January 17, 1984 123-140

No. 11. ROBERTS, TYSON R., AND MAURICE KOTTELAT. Description and osteology of
Thryssocypris, a new genus of anchovylike cyprinid fishes, based on two new
species from Southeast Asia. Published January 17, 1984 141-158

No. 12. KAVANAUGH, DAVID H. Studies on Nebriini (Coleoptera: Carabidae), V. New

Nearctic Nebria taxa and changes in nomenclature. Published July 12, 1984... 159-177

No. 13. ROBERTS, TYSON R. Skeletal anatomy and classification of the neotenic Asian
salmoniform superfamily Salangoidea (icefishes or needlefishes). Published July
12, 1984 179-220

No. 14. TRICAS, TIMOTHY C., AND JOHN E. McCosKER. Predatory behavior of the white
shark (Carcharodon carcharias), with notes on its biology. Published July 12,
1984 221-238

No. 1 5. NEWBERRY, ANDREW TODD. Dendrodoa (Styelopsis) abbotti, sp. nov. (Styelidae,
Ascidiacea) from the Pacific Coast of the United States, and its impact on some
gonadal criteria of its genus and subgenus. Published September 19,
1984 239-248

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Pages

No. 1 6. MCMILLAN, CHARMION B., AND ROBERT L. WISNER. Three new species of seven-
gilled hagfishes (Myxinidae, Eptatretus) from the Pacific Ocean. Published De-
cember 11, 1984 249-267

No. 17. ALMEDA, FRANK. New and noteworthy additions to the Melastomataceae of

Panama. Published December 11, 1984 269-282

No. 18. COMPAGNO, LEONARD J. V., AND TYSON R. ROBERTS. Marine and freshwater
stingrays (Dasyatidae) of West Africa, with description of a new species. Pub-
lished December 1 1 , 1 984 283-300

No. 19. FITCH, JOHN E., AND STEPHEN J. CROOKE. Revision of Eastern Pacific catalufas
(Pisces: Priacanthidae) with description of a new genus and discussion of the
fossil record. Published December 11, 1984 30 1-3 1 5

No. 20. ROBERTS, TYSON R. Amazonsprattus scintilla, new genus and species from the
Rio Negro, Brazil, the smallest known clupeomorph fish. Published December
11,1984 3 1 7-32 1

Index to Volume 43 323-329

Instructions to Authors .... 331-332

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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 1, pp. 1-16, 10 figs.

June 15, 1982

By
Tyson R. Roberts

California Academy of Sciences, Golden Gate Park, San Francisco, California 94118

ABSTRACT: The tetraodontid pufferfish genus Chonerhinos, restricted to fresh water in Southeast Asia,
comprises five species, four of which are described as new. The species differ in adult size, coloration, orientation
of squamation, depth of caudal peduncle, size of nasal organ, food habits, and geographical distribution. The
most widely distributed, C. nefastus n.sp., occurs in southern, western, and northern Borneo, the Malay
Peninsula, Thailand, Vietnam, Kampuchea, and Laos; it feeds mainly on fish fin rays and scales, and has a
slender caudal peduncle and the smallest nasal organ. Chonerhinos modestus (Bleeker, 1850), in western Borneo
and Sumatra, with perhaps the most varied diet, is the largest species and has the deepest caudal peduncle.
The distinctively colored C. amabiUs n.sp., with the largest nasal organ, occurs in western Borneo and Sumatra
and feeds almost exclusively on large aquatic insects. The two new species C. silus, with a moderately deep
caudal peduncle, and C. remotus, with a slender caudal peduncle, have varied diets including insects, and are
known only from northern and northeastern Borneo.

INTRODUCTION

The freshwater pufferfish genus Chonerhinos
currently includes a single species, C. modestus
(Bleeker, 1850), reported from localities
throughout much of Southeast Asia. The nomi-
nal species C. africanus Boulenger, 1909,
known only from the holotype supposedly col-
lected in the interior of the Congo basin, has
been identified as a junior synonym of C. mo-
destus with incorrect locality data (Roberts
1981; herein). The species formerly known as C.
naritus (Richardson, 1848), from marine, brack-
ish, and perhaps freshwater habitats along the
coasts of the South China Sea and eastern Indian
Ocean, has been placed in a monotypic genus,
Xenopterus (Fraser-Bruner 1943; Tyler 1980;
herein).

I undertook this revision because three
species of Chonerhinos were obtained during

my ichthyological survey of the Kapuas basin in
western Borneo (Kalimantan Barat, Indonesia)
in 1976.

MATERIAL EXAMINED AND METHODS

More than 250 specimens of Chonerhinos
from throughout the range of the genus were
examined during this study. These are deposited
in the British Museum (Natural History), Lon-
don, BMNH; California Academy of Sciences,
San Francisco (CAS), including material for-
merly deposited at Stanford University, Stan-
ford (SU); Field Museum of Natural History,
Chicago (FMNH); Museum Geneve, Geneva
(MG); Museum National d'Histoire Naturelle,
Paris (MNHN); Museum Zoologicum Bogo-
rense, Bogor, Indonesia (MZB); Musee Royal
de 1'Afrique Centrale, Tervuren (MRAC); Nat-
ural History Museum, Basel (NHMB); Rijks-

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1

museum van Natuurlijke Historic, Leiden
(RMNH); University of Michigan Museum of
Zoology, Ann Arbor (UMMZ); U.S. National
Museum of Natural History, Smithsonian Insti-
tution, Washington, D.C. (USNM); and Zoolog-
ical Museum, Universiteit van Amsterdam, Am-
sterdam (ZMA).

Length of specimens is given as standard
length unless total length is expressly indicated,
and all proportional measurements are given as
times in standard length (SL). Vertebral counts
were made from radiographs prepared in the
Department of Ichthyology, CAS. Sections of
skin anterior and ventral to the pectoral fin were
removed with a scalpel and stained in alizarin to
facilitate illustration of the scales. Orientation of
the scales is also obvious in radiographs and can
be observed in whole specimens without special
preparation.

Chonerhinos Bleeker

Chonerhinos BLEEKER, 1 854: 259-260 (type-species Tetraodon
modest us Bleeker, 1850, by subsequent designation of Fra-
ser-Bruner 1943:16).

Chonerhinus BLEEKER, 1865:213 (unjustified spelling change).

DESIGNATION OF TYPE-SPECIES. — Fraser-
Bruner (1943) is apparently the first author to
have properly designated a type-species for
Chonerhinos. The original description of the ge-
nus is as follows: "Chonerhinos Blkr [is
gekenmerkt] door trechtervormige verdieping
ter plaatse der neusopeningen met verhevene
randen, lange rug- en aarsvinnen, zigtbare zijlijn
en onegekielden rug . . . van Chonerhinos 2 t.
w. Chonerhinos modestus Blkr = Tetraodon
modestus Blkr olim (van Borneo, Sumatra),
Chonerhinos naritus Blkr = Tetraodon naritus
Richds (van Borneo)/' Thus, Bleeker included
two species in his original account of Chone-
rhinos and did not indicate a type-species. Hoi-
lard (1857) defined Xenopterus (type-species X.
belle ngeri = X. naritus, by monotypy) in such
a way that it excludes Chonerhinos, which,
however, he did not mention by name. Gill
(1892) discussed the nomenclatural history of
Chonerhinos (and Xenopterus} at length but
oddly did not mention the lack of a type-species.
Jordan (1919:256) incorrectly stated that Tetrao-
don modestus Bleeker is the "orthotype" of
Chonerhinos, meaning that Bleeker (1854) indi-
cated or distinctly implied that this species is the
type-species.

DIAGNOSIS. — Chonerhinos and its close rela-
tive Xenopterus differ from all other tetraodon-
tids in having three lateral line canals on side of
body instead of one, two, or none; dorsal fin
with 22 or more rays; anal fin with 18 or more
rays; at least 24 vertebrae; and prefrontal bones
absent (Tyler, 1980). Chonerhinos differs from
Xenopterus in its smaller adult size, less exten-
sive squamation, less exposed olfactory lamel-
lae, and fewer fin rays and vertebrae. The largest
Chonerhinos I have examined is 106 mm; Xen-
opterus attains at least twice this size. In Cho-
nerhinos the scales are relatively small and re-
stricted to the head and body ventral to the level
of the pectoral fin; in Xenopterus the scales are
relatively large and extend dorsally to the pec-
toral fin. In Chonerhinos the olfactory lamellae
are largely covered by nasal flaps in broad con-
tact; in Xenopterus the nasal flaps are greatly
reduced and the olfactory lamellae are conse-
quently almost entirely exposed. Chonerhinos
has 22-28 dorsal-fin rays, 18-22 anal-fin rays,
13-17 pectoral-fin rays, and 24-28 vertebrae; the
same counts in Xenopterus are 32-38, 28-29,
18-19, and 29-30.

REMARKS.— Tyler (1980) stated that Chone-
rhinos and Xenopterus are highly specialized
tetraodontids which have secondarily increased
the number of dorsal- and anal-fin rays and ver-
tebrae, elaborated the lateral line system, in-
creased the number and size of the olfactory la-
mellae, and increased the size of at least some
of the scales; and that the greater numbers of
vertebrae and fin rays in Xenopterus as well as
the structure of the skull indicates that it is the
more specialized of the two. In Chonerhinos,
according to Tyler, apart from the absence of
the prefrontal bones, the skull is not markedly
different from that in many species of the tet-
raodontid genera Monotreta, Chelonodon, and
Tetraodon, whereas in Xenopterus the frontals
are much more laterally expanded and thickened
than in Chonerhinos, forming a large plate over
most of the dorsal surface of the skull, and the
supraoccipital crest is wider and heavier; in
large specimens the two frontals may become
indistinguishably fused to each other in the mid-
dle of their lengths (Tyler 1980:340, fig. 274). I
have examined two X. naritus from Sarawak,
BMNH 1894.1.19.86-87, 71.2 and 108 mm. Ra-
diographs reveal that the frontal bones, supra-
occipital crest, supraneural bone, anteriormost

ROBERTS: FRESHWATER PUFFERFISH

FIGURE 1. Scales on side of body immediately anterior and ventral to pectoral fin (each square = 5x5 mm): (a) Xenopterus
naritus, 71.2 mm, BMNH 1894.1.19.86; (b) Chonerhinos modestus, 48.4 mm, USNM uncatalogued; (c) Chonerhinos silus, 48.5
mm, FMNH 68815; (d) Chonerhinos remotus, 49.9 mm, FMNH 68475; (e) Chonerhinos nefastus, 48.3 mm, CAS 49507; (/)
Chonerhinos amabilis, 48.7 mm, MZB 3973.

anal-fin pterygiophore, and posteriormost neural
and haemal spines are enormously thickened or
hypertrophied, far out of proportion to neigh-
boring bony elements. They appear to be hy-
perosteotic (and in the case of the frontal bones,
partially synosteotic), and therefore, I am du-
bious about their phylogenetic significance and
their being used as characters to distinguish
Xenopterus from Chonerhinos. Other differ-
ences between the two genera, cited above and
in Tyler (1980), are sufficient to merit their sep-
aration.

Chonerhinos is known only from fresh water.
Xenopterus, so far as I have been able to deter-
mine, is marine or estuarine. There do not seem
to be any museum specimens of Xenopterus
with locality data from fresh water, and state-
ments in the literature that Xenopterus occurs
in fresh water (e.g., Cantor 1850:384; Weber and
de Beaufort 1962:373) appear to be based at least
partly on misinformation or confusion with Cho-
nerhinos.

In Chonerhinos and Xenopterus, as in many
other tetraodontids, each scale has a spinelike

distal portion which projects more or less
straight out from the skin when erected, as usu-
ally occurs when the fish inflates itself. When
the scales are not erect, they are partially or
wholly retracted beneath the skin, and the
spines may be oriented dorsally, dorsoposte-
riorly, or posteriorly, depending upon the
species (Fig. 1).

Size and shape of the jaw-teeth appear to be
nearly identical in all species of Chonerhinos.
One or two specimens of each species were dis-
sected to permit observation of the gill rakers;
all of the species have about 8-10 total gill rakers
on each gill arch (sometimes fewer on the first
arch). I have not attempted to distinguish the
species by differences in the pathways of the
lateral line canals. These are difficult to observe
in many specimens, and they seem to be highly
variable among individual specimens, often
being irregularly interrupted or running into
each other (Tyler 1980:fig. 223) and frequently
differing in their courses on opposite sides of a
specimen. Neither have I attempted to distin-
guish the species by counts of olfactory lamel-

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1

TABLE 1. PROPORTIONAL MEASUREMENTS IN Chonerhinos (expressed as times in standard length).

C. amabilis

C. modestus

C. nefastus

C. remotus

C. silus

SL mm

35.6-70.4

46.8-106

19.0-70.7

32.8-61.4

32.7-81.8

Eye

7.8-11.2

9.4-14.1

7.2-11.7

8.6-12.0

8.8-12.9

Nasal organ length

10.1-17.9

14.7-25.3

17.3-27.4

12.5-20.1

11.2-20.0

Snout length

6.0-7.2

6.4-7.5

5.8-7.1

6.6-8.7

6.3-8.1

Interorbital width

5.1-6.1

4.6-6.8

4.5-7.0

5.2-7.0

4.9-6.6

Pectoral-fin base length

10.5-12.2

9.5-11.5

9.9-13.5

8.9-11.1

9.1-12.4

Caudal peduncle depth

7.3-S.3

6.7-7.4

7.8-9.9

7.6-9.9

7.1-8.2

Caudal peduncle length

5.0-6.5

5.2-6.4

4.4-5.9

4.6-6.3

4.6-6.5

lae, the number of which seems to be highly
variable within each species, as is the size of the
nasal organ itself (Table 1).

PROPORTIONAL MEASUREMENTS;
MERISTIC FEATURES

Proportional measurements, in most instances
broadly overlapping and of little help in distin-
guishing species, are presented in Table 1. Fre-
quencies of counts of fin rays and vertebrae,
diagnostic for the genus but differing slightly
among species and of little or no help in identi-
fying individual specimens, are presented in Ta-
bles 2-3. Except in a few instances when counts
or measurements are particularly useful for def-
inition of species, these data are not repeated in
the text.

KEY TO SPECIES OF Chonerhinos

la. Scales on side of body anterior and ven-
tral to pectoral fin with spines directed
posteriorly (Fig. le-f) 2

Ib. Scales on side of body anterior and ven-
tral to pectoral fin with spines directed
dorsally or dorsoposteriorly (Fig. \b-d)

_ 3

2a. A roundish dark spot in middle of caudal
peduncle; dorsal and anal fins always with
angulated margins; upper lip not project-
ing beyond lower lip; exposed portion of
eye round; nasal organ relatively large, its

length 10. 1-17. 9 (times in SL)

C. amabilis

2b. No spot on caudal peduncle; dorsal and
anal fins usually with rounded margins;
upper lip usually projecting beyond lower
lip; exposed portion of eye usually hori-
zontally oval, especially in larger speci-
mens; nasal organ relatively small, its
length 17.3-27.4 C. nefastus

3a. Depth of caudal peduncle 6.7-7.4; upper
and lower lips about equally projecting or
lower lip slightly protruding; snout gently
sloping; scales on side of body anterior
and ventral to pectoral fin, very close-set
with spines directed dorsally (Fig. Ib);
anal-fin rays 20-22, modally 22 (Table 2);

adult size to 106 mm

C. modestus

3b. Depth of caudal peduncle 7.2-9.9; lower
lip usually projecting beyond upper lip;
snout strongly sloping; scales on side of

TABLE 2. FREQUENCIES OF FIN RAY COUNTS IN Chonerhinos.

Dorsal fin

Anal fin

Pectoral fin

18 19

13 14

C. amabilis -

_ _

C. modestus -

C. nefastus -

5 28

C. remotus 5

6 41

C. silus -

1 2

_ 2

ROBERTS: FRESHWATER PUFFERFISH

FIGURE 2. Chonerhinos amabilis, 45.2 mm, MZB 3972 (holotype).

body anterior and ventral to pectoral fin
not as close-set and with spines directed
dorsoposteriorly (Fig. \c-d); anal-fin rays
18-22, rarely 22, modally 19 or 20 (Table
2); adult size to 82 mm 4

4a. Caudal peduncle moderately deep, its
depth 7.2-8.2; dorsal-fin rays 23-28, av-
erage 25.4 C. silus

4b. Caudal peduncle slender, its depth 7.6-9.9;

dorsal-fin rays 22-26, average 23.6

__ C. remotus

Chonerhinos amabilis new species

(Figure 2)

Chonerhinus naritus WEBER AND DE BEAUFORT, 1962:374
(specimens reported from "Labang hara, soengei Serawai").

Chonerhinus modestus WEBER AND DE BEAUFORT, 1962:fig.
84.

HOLOTYPE. — MZB 3972, 45.2 mm, Kapuas R. 6 km w of
Putussibau, Kapuas Ichthyological Survey, 9 Aug. 1976.

PARATYPES. — CAS 49504, 45.0 mm, same data as holotype;
MZB 3973, 48.7 mm, Kapuas basin, Sungai Landok at Nga-
bang, 83 km ENE of Pontianak, Kapuas Ichthyological Survey,
15 July 1976; MZB 3974, 41.8 mm, Kapuas basin, Sungai Pi-
noh 20-60 km upstream from Nangapinoh, Kapuas Ichthyo-

TABLE 3. FREQUENCIES OF VERTEBRAL COUNTS IN Chonerhinos.

C. amabilis

C. modestus C.

nefastus

C. remotus

. silus

= 24(1)

9? + 16 = 25? (1)

= 25 (2)

= 25 (1)

9? +

= 25? (1)

= 25 (3)

10 +

= 25 (2)

9 + 17 = 26(1)

= 26 (1)

10 + 16 = 26 (9)

10 + 16 = 26 (3) 10

= 26(1)

= 26 (8)

10 +

= 26 (8)

= 27 (6)

10 +

= 27 (1)

11? + 16 = 27? (1)

11?

= 27? (2)

11? +

= 271(1)

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1

FIGURE 3. Chonerhinos modestus: (a) as illustrated in Bleeker 1865; length, locality, and present disposition of specimen
unknown; (b) 78.7 mm, RMNH 26931 (neotype).

logical Survey, 22-26 July 1976; MZB 3975, 38.3 mm, Kapuas
R. near Kampong Nibung, 7 km NE of Selimbau, Kapuas Ich-
thyological Survey, 5-6 July 1976; MZB 3976 and USNM
230359, 2:35.9-36.8 mm, Kapuas R. 53 km w of Putussibau,
Kapuas Ichthyological Survey, 6-7 Aug. 1976; MZB 3977
and FMNH 94255, 2:35.6-46.0 mm, Kapuas R. about 23 km
wsw of Putussibau, Kapuas Ichthyological Survey, 8-9 Aug.
1976; MNHN 91.216, 36.9 mm, Kapuas basin, M. Chaper,
1890; RMNH uncat., 2:40.9-41.2 mm, Kapuas basin, Sintang,
July 1894; RMNH 7935, 4:55.5-68. 1 mm, Kapuas basin, Raun,
Mar.-May 1894; ZMA 108.912, 3:56.3-70.4 mm, Kapuas ba-
sin, Soengai Serawai, Lebang Hara, Witkamp, no date;
UMMZ 171708, 2:36.2-38.3 mm, Sumatra, Moesi R. at Moera
Klingi, A. Thienemann, 1913.

DIAGNOSIS. — Chonerhinos amabilis is readily
distinguished from all other members of the ge-

nus by its highly distinctive coloration, almost
all elements of which are visible in all specimens
examined, including some century-old speci-
mens which may have been dead for some time
before being preserved. These unique features
include a roundish dark spot in middle of caudal
peduncle, visible in all specimens; a large, dis-
tinctively shaped dark mark on dorsal surface of
head extending uninterrupted from just behind
upper lip to well behind the eyes, set off by pale
coloration on the upper lip, sides of snout, nasal
flaps, and skin dorsal to orbits; pale white or
milky coloration on ventral and lateral surfaces
of body extending very far dorsally; dark col-

ROBERTS: FRESHWATER PUFFERFISH

FIGURE 4. Chonerhinos modestus, 64.6 mm, CAS 49505.

oration on dorsal surface of body markedly en-
hanced around base of dorsal fin; and a small
dark or dusky oval spot with indistinct margins
near tip of chin (very faint or absent in some
specimens). In addition, C. amabilis tends to
have the largest nasal organ of any Chonerhi-
nos, and thus of any tetraodontid (Tyler
1980:290); relatively large dorsal and anal fins
with angulated (rather than rounded) margins;
and scales on side of body anterior and ventral
to pectoral fin relatively small, few in number,
and with spines directed posteriorly (Fig. If).
ETYMOLOGY. — Latin amabilis, lovely.

Chonerhinos modestus (Bleeker)

(Figures 3-5)

Tetraodon (Arothron) modestus BLEEKER, 1850:16 (type-lo-
cality "Banjermassing, in fluviis").

Chonerhinos modestus BLEEKER, 1854:260.

Chonerhinus africanus BOULENGER, 1909:201 (type-locality
"riv. Sankuru. a Kondue Kasai, Congo").

NEOTYPE.— RMNH 26931, 78.7 mm, Kapuas basin, Sang-
gau, Westenenk, 1894.

ADDITIONAL MATERIAL EXAMINED. — RMNH uncat.,

2:49.2-59.2 mm, same data as neotype; RMNH 7934,
3:50.0-58.9 mm, Kapuas basin, Sintang, July 1894; CAS 49505
and MZB 3978, 2:64.6-106 mm, Kapuas R. about 23 km wsw
of Putussibau, Kapuas Ichthyological Survey, 8-9 Aug. 1976;
MZB 3979 and USNM 230360, 2:46.8-48.4 mm, Kapuas
R. at Silat, Kapuas Ichthyological Survey, 17 Aug. 1976;
BMNH 1846.6.22.75, 86.1 mm, Borneo, Frank Collection, no
date; BMNH 1867.11.28.125, 87.3 mm, Borneo, Bleeker Col-
lection, no date; RMNH 12004, 3:66.6-81.1 mm, Sumatra,
Lahat, Bleeker Collection, 1850-60; NHMB 822-824,
3:44.7-73.5 mm, Sumatra, Indragiri, H. A. von Meckel, 1895;
RMNH 7344 (part only), 8:47.9-62.0 mm, no locality data,
Bleeker Collection, no date; MRAC 15306, 52.5 mm, "Congo,
Sankuru River, Kasai" (holotype of C. africanus).

SELECTION OF NEOTYPE. — Identification of
C. modestus presented a difficult and taxonom-
ically important problem which I have resolved
by selecting a neotype. The holotype is lost or
at least it cannot be positively identified, and the
original description fits all five species of Cho-
nerhinos about equally well. In order to facilitate
the following discussion the original description
(Bleeker 1850:16) is reproduced here in its en-
tirety:

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1

FIGURE 5. Chonerhinos modestus: (a) 52.5 mm, MRAC 15306 (holotype of C. africanus); (b) 48.4 mm, USNM 230360.

Tetraodon (Arothron) modestus Blkr.

Tetraod. corpore oblongo compresso, altitudine 4 circiter
in ejus longitudine, latitudine 2 in altitudine; vertice, dorso,
lateribus caudaque laevibus, pectore genisque scabris; ca-
pite obtuso; lineo rostro-dorsali convexa; maxilla superiore
paulo prominente; oculis paulo superis; tentaculis nasalibus
2 conicis obtusis loco narium; linea lateral! inconspicua;
sacco pneumatico parvo; ano ante pinnam dorsalem sito;
pinnis dorsali et anali obtusis angulatis angulis rotundatis,
pectoralibus emarginatis, caudali truncata vel leviter emar-
ginata 5 in longitudine corporis; colore corpore supra viridi
infra argenteo, pinnis hyalino-viridescente.

D. 5/20. P. 2/12. A. 3/20. C. 9 vel 11 et lat. brev.

Habit. Banjermassing, in fluviis.

Longitude speciminis unici 60'".

Bleeker almost invariably recorded the length of
his specimens as total length in millimeters
(pers. commun. M. Boeseman, RMNH). Thus,
the last two lines of the description indicate that

it was based on a single specimen, the holotype,
total length 60 mm, from riverine habitat at
Bandjarmasin, southeastern Borneo (Barito ba-
sin). Bleeker obtained in all 59 specimens which
he identified as C. modestus, total lengths
46-126 mm, from Palembang (=Lahat?), Su-
matra, and Sambas, Pontianak, and Sintang as
well as Bandjarmasin in Borneo (Bleeker
1865:78). All extant "C. modestus" from Bleek-
er's collection apparently are deposited in the
BMNH, RMNH, and ZMA. The BMNH and
ZMA each have a single Bleeker specimen, both
of which are too large to be the holotype:
BMNH 1867.11.28.125, 87.3 mm, Borneo, ex-
amined by me, and ZMA 102.263, 104 mm, Bor-
neo, examined for me by H. Nijssen. The
RMNH has two lots, RMNH 12004, 3:66.6-81.1

ROBERTS: FRESHWATER PUFFERFISH

mm, Sumatra, Lahat ( = Palembang?), and
RMNH 7344, 52:29.0-74.0 mm (total lengths
37-88 mm), without locality data. If the holotype
still exists, it presumably is in RMNH 7344.
Among the 52 specimens are 4 which approxi-
mate 60 mm in total length; thus, on the basis
of length alone, the holotype cannot be identi-
fied. Moreover, each of the four specimens dif-
fers by one or two fin rays in at least two of the
three counts reported by Bleeker for the dorsal,
anal, and pectoral fins of the holotype. In my
opinion, none of these specimens can reason-
ably be identified as the holotype, and since their
locality data are lost, a neotype should not be
selected from among them. Unfortunately, I
have been unable to find any specimen of Cho-
nerhinos with locality data from Bandjarmasin
or the Barito and do not know which of the
species occur(s) there.

As noted above, the original description of C.
modestus fits all five species of Chonerhinos
about equally well. All species of Chonerhinos
normally have 1 1 caudal-fin rays, and all species
are represented by specimens with 25 dorsal-fin
rays and 14 pectoral-fin rays. On the other hand,
none of the more than 250 specimens examined
have 23 anal-fin rays. The highest number of
anal-fin rays observed, 22, is usually found in
the species herein identified as C. modestus, but
also occurs in C. nefastus and C. situs. Color-
ation and its variation in the species of Chone-
rhinos are too poorly known at present to be of
much help in their identification, and Bleeker' s
description of coloration of the holotype cannot
be accepted without reservation since he did not
collect the specimen himself and could not have
observed it until it had been in preservative for
many days or weeks. Bleeker (1865:pl. 213, fig
8) published an excellent figure of a specimen
which he identified as C. modestus. The length,
locality, and date of collection of the specimen
figured are not recorded, but it is not the holo-
type. It is evidently a much larger specimen,
with lateral line canals on the body plainly vis-
ible, and differs also in fin-ray counts from the
holotype as described by Bleeker. I have not
tried to match up the figure with an extant spec-
imen, although it may well be part of RMNH
7344. The figure does, however, show a number
of features characteristic of the largest species
of Chonerhinos, with which I unhesitatingly
identify it. These features include its large size

(indicated by the large size of the published il-
lustration as well as by the relatively small eye);
scales with dorsally oriented spines; relatively
high counts of dorsal- and anal-fin rays; and
deep caudal peduncle. All four specimens of to-
tal length 60 mm in RMNH 7344 also belong to
this species. Thus, there is every reason to identi-
fy it as C. modestus, although we cannot be sure
that this is the same species obtained for Bleeker
at Bandjarmasin. In the absence of specimens
with locality data from Bandjarmasin or the Ba-
rito, a specimen from the Kapuas basin has been
selected as neotype. This specimen bears a
strong resemblance to Bleeker' s figure of C.
modestus (Figs. 3a-b).

DIAGNOSIS. — Chonerhinos modestus, attain-
ing at least 106 mm, apparently is the largest
species of Chonerhinos and has the deepest cau-
dal peduncle. Depth of caudal peduncle 6.7-7.4
(vs. 7.2-9.9 in all other Chonerhinos). Scales
relatively large and close-set, those on body an-
teroventral to pectoral fin with spines directed
dorsally, as in Xenopterus (vs. spines directed
dorsoposteriorly or posteriorly in all other Cho-
nerhinos). Upper and lower lips about equally
projecting or lower lip slightly protruding. Ex-
posed portion of eye round. Snout gently slop-
ing. Nasal organ moderately large, its length
14.7-25.3. Dorsal-fin rays 25-28; and anal-fin
rays 20-22 (generally fewer in other Chonerhi-
nos).

REMARKS ON SYNONYMY. — Most records of
C. modestus in the literature other than those
cited in the synonymy above refer in whole or
in part to other species of Chonerhinos.

Chonerhinos africanus was described briefly
(and without a figure) on the basis of a single
specimen supposedly obtained together with
other fish specimens by E. Luja in the Sankuru
River, Kasai, Congo basin, in 1908. No addi-
tional specimens of Chonerhinos have been
found in Africa, and the holotype has not been
compared previously to Chonerhinos from
Southeast Asia. I have examined the 52.5-mm
holotype (Fig. 5a), comparing it directly with
specimens of all five species of Chonerhinos,
and conclude that it is conspecific with C. mo-
destus. It has 26 dorsal-fin rays; 22 anal-fin rays;
15 pectoral-fin rays; 10+16 vertebrae; scales
relatively large, those on sides of body antero-
ventral to pectoral fin with spines directed dor-
sally; lower lip slightly protruding; snout gently

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1

FIGURE 5. Chonerhinos modestus: (a) 52.5 mm, MRAC 15306 (holotype of C. africanus); (b) 48.4 mm, USNM 230360.

Tetraodon (Arothron) modestus Blkr.

Tetraod. corpore oblongo compresso, altitudine 4 circiter
in ejus longitudine, latitudine 2 in altitudine; vertice, dorso,
lateribus caudaque laevibus, pectore genisque scabris; ca-
pite obtuso; lineo rostro-dorsali convexa; maxilla superiore
paulo prominente; oculis paulo superis; tentaculis nasalibus
2 conicis obtusis loco narium; linea lateral! inconspicua;
sacco pneumatico parvo; ano ante pinnam dorsalem sito;
pinnis dorsal! et anali obtusis angulatis angulis rotundatis,
pectoralibus emarginatis, caudali truncata vel leviter emar-
ginata 5 in longitudine corporis; colore corpore supra viridi
infra argenteo, pinnis hyalino-viridescente.

D. 5/20. P. 2/12. A. 3/20. C. 9 vel 11 et lat. brev.

Habit. Banjermassing, in fluviis.

Longitude speciminis unici 60'".

Bleeker almost invariably recorded the length of
his specimens as total length in millimeters
(pers. commun. M. Boeseman, RMNH). Thus,
the last two lines of the description indicate that

ROBERTS: FRESHWATER PUFFERFISH

As noted above, the original description of C.
modestus fits all five species of Chonerhinos
about equally well. All species of Chonerhinos
normally have 1 1 caudal-fin rays, and all species
are represented by specimens with 25 dorsal-fin
rays and 14 pectoral-fin rays. On the other hand,
none of the more than 250 specimens examined
have 23 anal-fin rays. The highest number of
anal-fin rays observed, 22, is usually found in
the species herein identified as C. modestus, but
also occurs in C. nefastus and C. silus. Color-
ation and its variation in the species of Chone-
rhinos are too poorly known at present to be of
much help in their identification, and Bleeker' s
description of coloration of the holotype cannot
be accepted without reservation since he did not
collect the specimen himself and could not have
observed it until it had been in preservative for
many days or weeks. Bleeker (1865:pl. 213, fig
8) published an excellent figure of a specimen
which he identified as C. modestus. The length,
locality, and date of collection of the specimen
figured are not recorded, but it is not the holo-
type. It is evidently a much larger specimen,
with lateral line canals on the body plainly vis-
ible, and differs also in fin-ray counts from the
holotype as described by Bleeker. I have not
tried to match up the figure with an extant spec-
imen, although it may well be part of RMNH
7344. The figure does, however, show a number
of features characteristic of the largest species
of Chonerhinos, with which I unhesitatingly
identify it. These features include its large size

REMARKS ON SYNONYMY. — Most records of
C. modestus in the literature other than those
cited in the synonymy above refer in whole or
in part to other species of Chonerhinos.

Chonerhinos africanus was described briefly
(and without a figure) on the basis of a single
specimen supposedly obtained together with
other fish specimens by E. Luja in the Sankuru
River, Kasai, Congo basin, in 1908. No addi-
tional specimens of Chonerhinos have been
found in Africa, and the holotype has not been
compared previously to Chonerhinos from
Southeast Asia. I have examined the 52.5-mm
holotype (Fig. 5a), comparing it directly with
specimens of all five species of Chonerhinos,
and conclude that it is conspecific with C. mo-
destus. It has 26 dorsal-fin rays; 22 anal-fin rays;
15 pectoral-fin rays; 10 + 16 vertebrae; scales
relatively large, those on sides of body antero-
ventral to pectoral fin with spines directed dor-
sally; lower lip slightly protruding; snout gently

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1

FIGURE 6. Chonerhinos nefastus, 47.0 mm, MZB 3980 (holotype).

sloping; eye 10.7; olfactory organ 18.7; snout
7.2; interorbital width 5.25; depth caudal pedun-
cle 7.4; length caudal peduncle 5.9; and pectoral-
fin base 10.8. The caudal peduncle depth and
anal-fin ray count, while not conclusive, agree
best with C. modestus. Presumably, the speci-
men originated somewhere in Southeast Asia
and somehow became mixed with Luja's mate-
rial when it was sent on loan to the BMNH for
Boulenger to study.

Chonerhinos nefastus new species

(Figure 6)

Chonerhinos modestus D'AUBENTON AND BLANC, 1966:561
(Mekong basin, Kampuchea); TAKI 1974:199-200, fig. 187
(Mekong basin, Laos); IMAKI et al. 1978:29, pi. 18 (Kapuas
R. at Sintang); TYLER 1980 (fig. 223?).

HOLOTYPE.— MZB 3980, 47.0 mm, Kapuas R. 29 km w of
Putussibau, Kapuas Ichthyological Survey, 11 Aug. 1976.

PARATYPES.— CAS 49506 and MZB 3981, 4:34.3^3.3 mm,
Kapuas R. near Kampong Nibung, 7 km NE of Selimbau, Ka-
puas Ichthyological Survey, 5-6 July 1976; BMNH 1982.3.29.
254-5 and MZB 3982, 3:36.6-43.7 mm, Kapuas R. 53 km w
of Putussibau, Kapuas Ichthyological Survey 6-7 Aug. 1976;
IRSNB 632, MZB 3983, ROM 38601, and USNM 230361,
6:32.9-60.2 mm, Kapuas R. about 23 km wsw of Putussibau,
Kapuas Ichthyological Survey, 8-9 Aug. 1976; MZB 3984,

51.7 mm, Kapuas basin, small tributary of Sungai Mandai 17
km wsw of Putussibau, Kapuas Ichthyological Survey, 10
Aug. 1976; MZB 3985, 64.9 mm, Kapuas basin, Sungai Mandai
Kechil, 18 km wsw of Putussibau, Kapuas Ichthyological Sur-
vey, 11 Aug. 1976; CAS 49507 and MZB 3986, 3:36.7-57.8
mm, Kapuas basin, Sungai Tawang near Danau Pengembung,
Kapuas Ichthyological Survey, 14-15 Aug. 1976; RMNH
7936, 61.8 mm, Kapuas basin, Sibau, June 1894; RMNH un-
cat., 25.5 mm, Kapuas basin, Sintang, July 1894; ZMA
1 10.220, 65.8 mm, Kapuas basin, Bunut, H. A. Lorentz, 26
June 1909; FMNH uncat. 3:38.2-50.1 mm, Sarawak, Niah R.,
T. Harrisson, 1 Apr. 1963; FMNH uncat., 2:64.8-70.7 mm,
Sarawak, Niah, T. Harrisson, no date; FMNH uncat.,
3:36.9-41.8 mm, Sarawak, Rejang basin, Baleh R. between
Sungai Mujong and Sungai Gaat, R. F. Inger, 3 Aug. 1956;
RMNH 7933, 2:56.6-68.0 mm, Mahakam basin, Tepoe, A. W.
Nieuwenhuis, 1896-97; MG 2058.94, 34.9 mm, Kalimantan
Tengah, Mentaya basin near Sampit, Pfeuffer, May 1980;
UMMZ uncat., 50.1 mm, Sumatra, Moesi R. at Moera Klingi,
A. Thienemann, 1913; SU 36040, 41.7 mm, Malay Peninsula, Per-
ak, Chandra dam, A. W. Herre, 18 Mar. 1923; UMMZ 197038,
43.7 mm, Thailand, Songkhla Lake off Patalung, K. F. Lagler,
6 Jan. 1965; UMMZ uncat., 48.0 mm, Thailand, Mekong ba-
sin, Ubon Ratchtani, Huay Phai, 16 Oct. 1975; UMMZ uncat.,
38.9 mm, Thailand, Mekong basin, Ubon Ratchtani, Huay
Kwang, 1 Oct. 1976; UMMZ uncat., 42.5 mm, Thailand, Me-
kong basin, Huay Kwang s of Khong Chiam, Arden, 7 Oct.
1975; UMMZ uncat., 30.9 mm, Thailand, Mekong basin, Mun
R. at Khong Chiam, Songrad and Buskirk, 19 July 1975;
UMMZ uncat., 3:15.4-32.2 mm, Thailand, Mekong R. and

ROBERTS: FRESHWATER PUFFERFISH

FIGURE 7. Chonerhinos remotus, 52.7 mm, FMNH 68475 (holotype).

tributaries from Ban Dan to Nakon Phanom, Mekong fish sur-
vey, Mar.-Apr. 1975; MNHN 1966.55-56, 9:21.6-48.1 mm,
Kampuchea, Mekong basin, Prek Tasom, F. d'Aubenton, 5
June and 9 Nov. 1961; MNHN 1966.57, 12:19.0-47.5 mm,
Kampuchea, Mekong basin, Prek Andor, F. d'Aubenton, 2
Dec. 1961.

DIAGNOSIS. — Chonerhinos nefastus differs
from all other species of Chonerhinos in having
upper lip usually projecting beyond lower lip;
nasal organ relatively small (Table 1); and ex-
posed portion of eyeball usually horizontally
oval rather than round or vertically oval. It dif-
fers from all other species except C. amabilis in
having scales on side of body anterior and ven-
tral to pectoral fin usually with spines directed
posteriorly (Fig. le), and from all except C. re-
motus in its slender caudal peduncle (Table 1).
Body usually without distinct color marks ex-
cept for a slightly darkened spot on dorsal sur-
face of head posterior to eyes.

COMMENTS. — The exposed portion of the
eyeball is distinctly horizontally oval in more
than half of the specimens examined. It is usu-
ally round in very small specimens, however,
and sometimes round in large specimens (in-
cluding the holotype). Most specimens have the
scales on the side of the body anterior and ven-
tral to the pectoral fin with the spines directed
posteriorly, as in Figure le. This character is
variable, however, and in a few specimens the
spines are directed posterodorsally or almost
dorsally. This is most noticeable in the sample
of 12 specimens from Prek Andor, 4 of which

have the spines more dorsally directed than is
usual in C. nefastus. The rest of the specimens
in the sample have the spines directed poste-
riorly or posterodorsally. Specimens from the
Mekong River differ from C. nefastus from oth-
er localities in having a dark transverse mark on
the dorsal surface of the snout between the up-
per lip and the nostrils.

ETYMOLOGY. — Latin nefastus, wicked,
abominable, in reference to the food habits (see
below).

Chonerhinos remotus new species

(Figure 7)

Chonerhinos modestus HERRE, 1940:55 (Sandakan District,
Sungei Segaliud and Sungei Sibuga); INGER AND CHIN
1962: 190-191, fig. 101 (Kinabatangan District).

HOLOTYPE. — FMNH 68476, 52.7 mm, Kinabatangan basin,
mouth of Sungai Deramakot, R. F. Inger and P. K. Chin, 27
Apr. 1956.

PARATYPES. — FMNH uncat., 9:32.8-54.4 mm, same data as
holotype; CAS 49743 and FMNH 68475, 61:29.1-56.8 mm,
Kinabatangan R. below mouth of Malubok R., R. F. Inger and
P. K. Chin, 25 Apr. 1956; FMNH 68474, 3:47.2-54.4 mm,
Kinabatangan R. at Deramakot camp, R. F. Inger and P. K.
Chin, 24 Apr. 1956; FMNH 44931, 38.3 mm, Kinabatangan
District, N. Borneo Fisheries Dept., 20 Jan. 1949; SU 33487,
2:60.5-61.4 mm, Sandakan District, Sibugal R. (=Sungai Si-
buga), A. W. Herre, 19 Apr. 1938; SU 33563, 10:30.5-40.4
mm, Sandakan District, Segaliud R., A. W. Herre, 4 Feb.
1937.

DIAGNOSIS. — Chonerhinos remotus is most
similar to C. silus, from which it differs in hav-
ing a more slender caudal peduncle (Table 1);
fewer dorsal- and anal-fin rays on the average

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1

FIGURE 8. Chonerhinos silus, 44.9 mm, FMNH 68477 (holotype).

(Table 2); an even more strongly sloping snout;
lower lip almost always strongly projecting be-
yond upper lip (vs. lower lip slightly projecting
or equal to upper lip); and eye vertically oval or
round (vs. usually round). Scales anterior and
ventral to pectoral fin moderately large and
close-set, with spines directed posterodorsally
(Fig. \d). No distinctive color marks except for
a well-defined dark blotch on dorsal surface of
head posterior to eyes.

ETYMOLOGY. — Latin remotus, remote, in ref-
erence to the type-locality.

Chonerhinos silus new species
(Figure 8)

HOLOTYPE. — FMNH 68477, 44.9 mm, Sarawak, Rejang ba-
sin, Sungai Baleh between Sungai Mujong and Sungai Gaat,
R. F. Inger, 3 Aug. 1956.

PARATYPES.— CAS 49744, FMNH uncat., 36:35.0-60. 1 mm,
same data as holotype; FMNH 62987, 44.0 mm, Sarawak,
Niah R., Niah, Lord Medway, 22 Aug. 1959; FMNH 68813,
81.8 mm, Sarawak, Niah, T. Harrisson, no date; FMNH
68814; 2:44.1-68.7 mm, Sarawak, Niah, Niah R., Pengkalan
Lobang, T. Harrisson, 2-11 Nov. 1960; FMNH 68815,
16:37.1-72.9 mm, Sarawak, Niah R., T. Harrisson, 1 Apr.
1963; SU 33610, 32.7 mm, Sarawak, 16 miles [ca. 26 km] E of
Kuching, A. W. Herre, 16 Feb. 1937.

DIAGNOSIS. — Chonerhinos silus is most simi-
lar to C. remotus and C. modestus. Differences
between C. silus and C. remotus are set forth
above in the diagnosis of C. remotus. It differs
from C. modestus in attaining smaller adult size
(largest specimen examined 82 mm vs. 106 mm);
snout more strongly sloping; lips equally pro-

jecting, or lower lip variably protruding, fre-
quently much more so than in C. modestus;
scales anterior and ventral to pectoral fin with
spines projecting dorsoposteriorly (Fig. Ic) rath-
er than dorsally (Fig. \b); and caudal peduncle
relatively slender, its depth 7.2-8.2 (vs. 6.7-7.4).
C. silus tends to have fewer dorsal-, anal-, and
pectoral-fin rays than C. modestus (Table 2), but
the counts are broadly overlapping and of little
help in identifying individual specimens to
species.
ETYMOLOGY. — Latin silus, pugnosed.

COLORATION IN LIFE

Most of the specimens of Chonerhinos col-
lected during the 1976 Kapuas Ichthyological
Survey were caught at night and preserved be-
fore their coloration in life could be properly
observed. Colors of the 106-mm C. modestus,
gill-netted at night and removed the next morn-
ing, are recorded in my field notes and in a 35-
mm Kodachrome slide. It was pale blue dorsal-
ly, white on the sides and abdomen, and with a
reddish eye. It is my impression that the three
smaller C. modestus collected during the survey
were similarly colored. C. amabilis is described
in my field notes as lime-green dorsally, with a
darkened area along the base of the dorsal fin,
and a reddish eye; the round spot on the caudal
peduncle, so evident in preserved specimens,
was not observed during life (at least it is not
recorded in my field notes, and I do not recall

ROBERTS: FRESHWATER PUFFERFISH

having seen it in the live specimens). I suspect
that some C. amabilis were blue dorsally but
this is not recorded in my field notes. My
impression is that all C. nefastus caught during
the survey were pale green dorsally; at least this
was so in several specimens observed during the
day. I doubt that any of them were blue dorsally.
D'Aubenton and Blanc (1966) reported color-
ation of C. nefastus (as C. modestus) from the
Mekong basin in Kampuchea as green on the
back and white on the flanks and belly, while
Taki (1974) reported specimens from the Me-
kong in Laos as having "back and upper surface
of head and body olivaceous golden, underside
pale yellow to white. Dorsal and caudal fins
greenish yellow; anal fin pale yellow; pectoral
fins hyaline."

SEXES

Secondary sexual dimorphism is unknown in
Chonerhinos. I have examined ripe males and
gravid or ripening females in all five species.
Ovaries of the left and right sides are about
equally well developed. The following approxi-
mate counts of eggs and measurements of egg
diameters contained in the right ovary were
made; C. amabilis, 57.4 mm, 180 eggs, 1.1-1.9
mm; C. modestus, 106 mm, 800 eggs, 1.5-2.1
mm; C. nefastus, 56.5 mm, 100 eggs, 1.4-1.5
mm, 57.8 mm, 80 eggs, 1.3 mm, and 64.9 mm,
230 eggs, 1.3-1.6 mm; C. remotus, 54.2 mm, 85
eggs, 1.9-2.3 mm; and C. situs, 58.7 mm, 200
eggs, 1.5-2.1 mm. All of these specimens are
gravid except the three C. nefastus, which are
nearly ripe. In C. remotus I observed two gravid
females, 54.2 and 54.4 mm, and three spent fe-
males, 48.8, 51.4, and 52.7 mm (the holotype,
Fig. 7), with genitoanal areas much swollen.
Such swelling, perhaps present only in females
just before or after spawning, has not been ob-
served in other species.

FOOD HABITS

Food habits of Chonerhinos, determined by
complete or partial examination of gut contents
in more than 100 specimens, may be summa-
rized as follows: C. amabilis feeds almost ex-
clusively on large aquatic insects; C. modestus
feeds mainly on terrestrial insects, shrimps,
seeds, and to a less extent on whole fish, fin
rays, or scales; C. nefastus feeds mainly on fish
fin rays and scales, and to a lesser extent on

insects (aquatic and terrestrial); C. remotus and
C. silus feed mainly on insects aquatic and ter-
restrial), but also ingest vegetable matter and
other items. No fish remains were found in C.
amabilis, C. silus, or, excepting a single fish
scale in one specimen, C. remotus. Pieces of
clam flesh and gills were found in several C.
silus, and numerous small, whole clams in a sin-
gle C. nefastus, but otherwise molluscs were
absent. The food of the five species may be de-
scribed in more detail as follows.

In C. amabilis, 18 of 20 specimens contained
more or less abundant remains of insects, mainly
large aquatic forms; partial examination of the
gut contents of these specimens failed to reveal
any other food items. Of the remaining two
specimens, one contained moderate amounts of
an unidentified flocculent material, and one had
empty guts. This species is noteworthy in that
nearly all individuals had much food in their
guts, and in being the most stenophagic of any
species of Chonerhinos. In C. modestus, guts
were examined in 10 specimens, half of which
had empty guts. Of the remaining five, four con-
tained moderate to large amounts of insects
(mainly terrestrial), two had prawns, two had
seeds, two had fish scales, one had fish fin rays,
and one had the remains of a small whole cobitid
fish (identified by its Weberian apparatus). The
last C. modestus, the 106-mm specimen, is of
particular interest because of its large size and
because of the circumstances of its capture. It
was gill-netted together with a large catfish,
Pangasius polyuranodon (Fig. 9), which had
much of its abdominal wall and portions of its
anal and caudal fins and caudal peduncle bitten
away. I suspected that part of the damage may
have been done by the C. modestus, but careful
examination of its gut contents failed to reveal
any material from the Pangasius. While the C.
modestus may have regurgitated, its stomach
did contain other food items, and it seems more
likely that the Pangasius was ravaged by some
other predator, possibly C. nefastus. Of 31 C.
nefastus in which the gut contents were exam-
ined, 1 1 had more or less substantial amounts of
fish fin rays, six had fish scales, three contained
small pieces offish flesh, six had small to mod-
erate amounts of insects (terrestrial and aquat-
ic), two had unidentified debris or detritus, one
had numerous small, whole bivalves, and one
had a large amount of sand and grit; seven had

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1

FIGURE 9. An 106-mm Chonerhinos modestus gill-netted together with an 80-cm Pangasius polyuranodon catfish ravaged
by an unknown predator, possibly C. modestus or C. nefastus (Kapuas River near Putussibau).

empty guts. The Latin name nefastus refers to
the predominantly pterygophagous and lepido-
phagous habits of this species. Inger and Chin
(1962:191) reported gut contents of 11 C. re-
motus (as C. modestus) as follows: bits of leaves
(6); parts of terrestrial insects (6); Plecoptera
nymphs (3); Trichoptera larval cases (1); un-
identified insect larvae (3); Acarina (2); unspec-
ified parts of fishes (2). Of 21 C. remotus I ex-
amined, 18 had guts containing food items: 14
with insects (aquatic and terrestrial), 4 with
parts of higher plants, 1 with a mite, 1 with a
fish scale, and several with unidentified debris
or detritus. In 33 C. silus, 22 had guts containing
insects (aquatic and terrestrial), 6 contained
higher plant material (fine rootlets, leaf, seeds,
or seed pulp?), 1 had several pieces of a large,
spinulose oligochaete, and 1 had chunks of spiny
or hairy flesh (mammalian?); the remainder had
empty guts.

INTRASPECIFIC BITING

Intraspecific biting, although infrequently
documented, probably occurs in many members
of the family Tetraodontidae. In Fugu niphobles
(Jordan and Snyder, 1901), biting is an integral
part of spawning behavior: egg laying occurs on
the beach at high tide after a female has been
bitten on the sides by two to four males (Uno
1955). Many of the specimens of Chonerhinos
examined exhibited characteristically shaped
bite marks on the flanks and, even more fre-
quently, had portions of the median fins bitten
off. I suspect that much of the biting, at least in

C. nefastus, is inflicted by conspecifics. More
than half of the specimens examined of this
species had bite marks on the flanks or had por-
tions of the dorsal, anal, or caudal fins missing.
In many specimens these fins appear to have
been bitten repeatedly, as evidenced by scar tis-
sue and imperfect regeneration of fin rays. It is
noteworthy that this species feeds predominant-
ly upon fish fin rays (see above under Food Hab-
its). C. modestus and C. silus, both of which
occur sympatrically with C. nefastus, also ex-
hibit high frequencies of specimens with bite
marks and bitten fins, but it is unclear whether
this is a result of intraspecific attacks, attacks
by C. nefastus, or a combination of both. In all
three species the bite marks and fin damage ap-
pear to be about equally distributed between the
sexes, and between gravid and nongravid fe-
males. None of the specimens of C. amabilis
and C. remotus examined exhibited bite marks
on the flanks, and their fins were relatively un-
damaged, with little or no indication of fin-nip-
ping. Perhaps the generally pterygophagous and
lepidophagous feeding behavior of C. nefastus
was preceded by the evolution of an exception-
ally aggressive intraspecific biting and fin-nip-
ping behavior.

GEOGRAPHICAL DISTRIBUTION

Tetraodontidae is the only one of the nine
families of the large order Plectognathi or Te-
traodontiformes which has representatives that
occur in fresh water. About 25 of the approxi-
mately 140 described tetraodontid species are

ROBERTS: FRESHWATER PUFFERFISH

-30°

-20C

-10C

°C. amabilis
*C. modestus
•C. nefastus
*C. remotus
°C. silus

-oc

11(11

FIGURE 10. Geographical distribution of species of Chonerhinos.

endemic to fresh water. Carinotetraodon and
Chonerhinos, both from Southeast Asia, are the
only tetraodontid genera restricted to fresh
water. Other genera with freshwater species in-
clude Tetraodon or Monotreta in India, South-
east Asia, and New Guinea; Tetraodon in Afri-
ca; and Colomesus in South America. Two
features of the geographical distribution of
freshwater Tetraodontidae merit comment.
First, although marine tetraodontids extend into
high latitudes in the Northern and Southern
hemispheres, freshwater species occur only
within tropical latitudes. Second, the tropical
rivers with endemic tetraodontids generally
have rich ichthyofaunas dominated by primary
freshwater fishes.

Geographical distributions of the species of
Chonerhinos, based mainly on material exam-
ined in this study, are illustrated in Figure 10.
Two of the species, C. amabilis and C. modes-

tus, have distributions lying within the hydro-
graphic limits of the ancient Central Sundaland
River basin, now fragmented by the Java and
South China seas. I suspect that C. modestus
also occurs in Thailand but have not examined
specimens from there. The most widely distrib-
uted species, C. nefastus, occurs throughout the
area occupied by the Central Sundaland River
basin; it also occurs in northern and southern
Borneo and in the Mekong basin. Whether the
Mekong River once also formed part of the Cen-
tral Sundaland drainage is a matter under inves-
tigation. C. silus and C. remotus, in northern
and northeastern Borneo, have restricted distri-
butions entirely outside the limits of the Central
Sundaland drainage area. C. amabilis, C. mo-
destus, and C. nefastus occur sympatrically in
the Kapuas River and probably also in some
rivers in Sumatra including the Indragiri and

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1

Moesi. C. nefastus and C. silus occur sympatri-
cally in Sarawak (Rejang and Niah basins).

ACKNOWLEDGMENTS

It is a pleasure to thank the following individ-
uals for their help during this study: Oliver
Crimmen, Gordon Howes, and Alwyne Wheel-
er, BMNH; Marie-Louise Bauchot, Jaques Dag-
et, and Martine Desoutter, MNHN; Marinus
Boeseman, Peter van Helsdingen, and M. J. P.
van Oijen, RMNH; Han Nijssen, ZMA; Volker
Mahnert, MG; Donald J. Stewart, FMNH; Wal-
ter Rainboth, UMMZ; Dirk Thys van den Au-
denaerde, MRAC; Maurice Kottelat, Universite
de Neuchatel; and Lillian Dempster, Madeleine
Graham, W. I. Follett, Michael Hearne, and
James Jackson, CAS. Photography is by Al-
phonse Coleman, Museum of Comparative Zo-
ology, Harvard University, and Orrin Moon,
The Darkroom, San Rafael.

The ichthyological survey of the Kapuas basin
was sponsored by the Museum Zoologicum Bo-
gorense, Indonesian National Research Council,
and Smithsonian Tropical Research Institute.
Soetikno Woerjoatmodjo, Leo Poerwadi, and
Rajali assisted in the field. Research was done
during visits to the BMNH, MNHN, RMNH,
and ZMA, and at the California Academy of Sci-
ences and Tiburon Center for Environmental
studies, and was supported by National Science
Foundation grant DEB77-24759.

LITERATURE CITED

BLEEKER, P. 1850. Bijdrage tot de kennis der ichthyologische
fauna van Borneo, met beschrijving van 16 nieuwe soorten
van zoetwatervisschen. Nat. Tijdschr. Ned. Ind. 1:1-16.

. 1854. Vijfde bijdrage tot de kennis der ichthyolo-
gische fauna van Celebes. Nat. Tijdschr. Ned. Ind.
7:225-260.

. 1865. Atlas ichthyologique des Indes Orientales Neer-

landaises, vol. 5.

BOULENGER, G. A. 1909. Catalogue des poissons du Congo
du Musee d'Histoire naturelle de Luxembourg. Faune de
Sankuru a Kondue (Collection Ed. Luja). Monatsber. Ge-
sell. Luxemburg. Naturf., n. sen, 3:189-202.

CANTOR, T. E. 1850. Catalogue of Malayan fishes. J. Thomas,
Calcutta, xii + 461 p., 24 pis.

D'AUBENTON, F., AND M. BLANC. 1966. Poissons tetraodon-
tiformes du Cambodge. Bull. Mus. Natl. Hist. Nat. ser. 2,
38:554-561.

FRASER-BRUNER, A. 1943. Notes on plectognath fishes. —
VIII. The classification of the suborder Tetraodontoidea,
with a synopsis of the genera. Ann. Mag. Nat. Hist. ser. 11,
10:1-18.

GILL, T. N. 1892. Note on the genus Chonerhinos or Xe-
nopterus. Proc. U.S. Natl. Mus. 14:696-699.

HERRE, A. W. 1940. Additions to the fish fauna of Malaya
and notes on rare or little known Malayan and Bornean
fishes. Bull. Raffles Mus. 16:27-61.

HOLLARD, H. 1857. Etudes sur les Gymnodontes et en par-
ticulier sur leur osteologie et sur les indications qu'elle peut
fournir pour leur classification. Ann. Sci. Nat. (Paris), zool.,
ser. 4, 8:275-328.

IMAKI, A., A. KAWAMOTO, AND A. SUZUKI. 1978. A list of
freshwater fishes collected from the Kapuas River, West
Kalimantan, Indonesia. The Institute for Breeding Re-
search, Tokyo University of Agriculture, 50 p.

INGER, R. F., AND P. K. CHIN. 1962. The fresh-water fishes
of North Borneo. Fieldiana: Zool. 45: 1-263.

JORDAN, D. S. 1919. The genera of fishes, 2. Stanford Univ.
Publ., univ. ser., i-x + 163-284 + i-xiii p.

ROBERTS, T. R. 1981. Identification of the presumed African
freshwater fishes Micracanthus marchei (Belontiidae) and
Chonerhinos africanus (Tetraodontidae). Cybium, ser. 3,
5:91-92.

TAKI, Y. 1974. Fishes of the Lao Mekong basin. USAID
Mission to Laos, Agric. Div., vi + 232 p.

TYLER, J. C. 1980. Osteology, phylogeny, and higher classi-
fication of the fishes of the order Plectognathi (Tetraodon-
tiformes). NOAA Tech. Rep. NMFS Circ. 434, 422 p.

UNO, Y. 1955. Spawning habit and early development of a
puffer, Fugu (Torafugu) niphobles (Jordan et Snyder). J.
Tokyo Univ. Fish. 41:169-183.

WEBER, M., AND L. F. DE BEAUFORT. 1962. The fishes of
the Indo-Australian Archipelago, vol. 11. E. J. Brill, Lei-
den, ix + 481 p.

CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, California 94118

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 2, pp. 17-24, 7 figs. June 15' 1982

SYNONYMIES OF INDIAN OCEAN EELS, WITH THE

DESCRIPTION OF GYMNOTHORAX ENIGMATICUS,

A MORAY PREVIOUSLY KNOWN AS G. RUPPELI

By
John E. McCosker

Steinhart Aquarium, California Academy of Sciences, Golden Gate Park,
San Francisco, California 94118

and
John E. Randall

Bernice P. Bishop Museum, Honolulu, Hawaii 96818

ABSTRACT: The common, banded Indo-Pacific morays called Gymnothorax peteUi (Bleeker, 1856) and G.
ruppeli (McClelland, 1845) by recent authors are recognized as G. rueppelliae (McCleUand, 1845) and G.
enigmaticus n.sp., respectively. They are separable on the basis of coloration, vertebrae, and morphology and
have different geographic ranges. G. signffier Bliss, 1883, is placed in the synonymy of G. rueppelliae, along
with Muraena umbrofasciata Ruppeli, 1852; M. interrupta Kaup, 1856; Sideria chlevastes Jordan and Gilbert,
1883; G. leucacme Jenkins, 1904; and G. waialuae Snyder, 1904. The moray Uropterygius xanthopterus Bleeker,
1859, is recognized as distinct from V. marmoratus (Lacepede, 1803), and V. alboguttatus Smith, 1962, is
synonymous with it. Ophichthus retifer Fowler, 1935, from Durban, South Africa, is a synonym of (). erabo
(Jordan and Snyder, 1901), an ophichthid also known from Hawaii, Japan, and Taiwan.

INTRODUCTION the snout tip to the posterodorsal margin of the

In preparation for the publication of the eel gill opening; trunk length is taken from the end

section of the revised Sea Fishes of Southern of the head to mid-anus; maximum body depth

Africa (McCosker and Castle, MS), we assign does not include the median fins. Vertebral

several poorly known taxa to synonymy and counts (which include the hypural) were taken

provide a description for a common, conspicu- from radiographs. Materials used in this study

ously banded Indo-Pacific moray, Gymnothorax are housed at the following institutions: Acad-

ruppeli of earlier authors, which lacks a holo- emy of Natural Sciences of Philadelphia

type and scientific name. (ANSP); Bernice P. Bishop Museum (BPBM);

British Museum of Natural History (BMNH);

METHODS California Academy of Sciences (CAS); U.S.

Measurements are straight-line, made either National Museum of Natural History (USNM);

with a 300-mm ruler with 0.5-mm gradations (for Museum of Comparative Zoology, Harvard Uni-

total length, trunk length, and tail length) and versity (MCZ); J. L. B. Smith Institute of Ich-

recorded to the nearest 0.5 mm, or with dial cal- thyology, Rhodes University (RUSI); Natur-

ipers (all other measurements) and recorded to Museum Senckenberg (SMF); and the Scripps

the nearest 0. 1 mm. Body length comprises head Institution of Oceanography (SIO). Paratypes of

and trunk lengths. Head length is measured from the new species will also be deposited at the

[17]

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 2

FIGURE 1. Paratype of Gymnothorax enigmaticus n.sp., BPBM 9491, 335 mm TL, from Palau.

Museum National d'Histoire Naturelle, Paris
(MNHN), the BMNH, and the USNM.

FAMILY MURAENIDAE

Gymnothorax enigmaticus, new species
(Figure 1)

Holotype.— CAS 48815, 303.2 mm total length, a male (?)
collected with rotenone in a 0-1-m tidal flat, off southern cor-
ner Ngatchab Beach, Angaur I., Palau, Western Caroline Is.,
by H. DeWitt, Sumang, and Sengjch, 21 Oct. 1957.

Paratypes. — Collected in shallow coral reef flats and tide-
pools (0-3 m) using rotenone ichthyocides. PALAU: CAS 48823
(8; 51-299 mm), Angaur I., H. DeWitt et al., 22 Oct. 1957.
CAS 48817 (293 mm), Auluptagel I., H. Fehlmann et al., 10
Aug. 1955. CAS 48816 (335.5 mm), Ngethil I., Sumang and R.
Johaness, 13 July 1959. CAS 48822 (2; 238-257 mm), Aulong
I., Sumang, 5 Nov. 1959. CAS 48826 (301.5 mm), Urukthapel
I., H. Fehlmann et al., 19 Aug. 1955. BPBM 9491 (335 mm),
Malakal Harbor, A. Emery, 21 Apr. 1970. K.APINGAMARINGI:
CAS 48818 (163.8 mm), Thokotaman, R. Harry, 12 July 1954.
IFALUK ATOLL: CAS 48819 (174 mm), Falarik Islet, R. Harry,
26 Sep. 1953. GUAM: CAS 48820 (214.4 mm), N of Cocos Is.,
Nangauta and H. Fehlmann, 8 Oct. 1958. ENEWETAK ATOLL:

CAS 42377 (144 mm), Runit I., R. Nolan and L. Taylor, Jr.,
23 Feb. 1974. BPBM 8184 (127 mm), Enewetek I., J. Randall,
1 Dec. 1967. BPBM 22339 (2; 219-233 mm), Enjebi I., J. Ran-
dall et al., 27 Apr. 1978. BIKINI ATOLL: BPBM 12354 (310
mm), Eman I., V. Brock et al., 18 June 1947. LINE Is.: CAS
48825 (302 mm), Palmyra I., E. Herald et al., 16 Aug. 1951.
BPBM 7715 (2; 310-393 mm), Cooper I., J. Randall, 13 Nov.
1968. HONG KONG: CAS 48821 (3; 79-88 mm), Santa Cruz Is.,
Vanikoro I., R. Bolin, 30 Sep. 1958. INDONESIA: BPBM 20890
(2; 103-383 mm), Bali, Sanur Beach, J. Randall, 18 July 1977.
THAILAND: BPBM 22827 (460 mm), Similan I., Ko Miang, J.
Randall, 14 Feb. 1979. PHILIPPINES: CAS 48824 (2; 508-518.5
mm), Negros Oriental, D. Empero, 28 July 1958.

DIAGNOSIS. — A moderate-length species of
Gymnothorax with anus before midbody; tubu-
lar anterior nostrils; uniserial jaw and vomerine
teeth; and cream body coloration with 17-21 dis-
tinctive brown bands encircling head and body
and extending onto fins.

DESCRIPTION OF HOLOTYPE (followed paren-
thetically by mean and range of the condition of
holotype and nine paratypes). — Greatest depth

McCOSKER & RANDALL: INDIAN OCEAN EELS

FIGURE 2. Gymnothorax rueppelliae, BPBM 18412, 339 mm TL, from Enewetak.

of body 16.8 (19.0; 15.4-22.7) times in total
length (TL). Tail longer than body, its length
1.76 (1.76; 1.71-1.82) in TL. Head 7.94 (7.69;
7.19-8.19) and trunk 3.26 (3.30; 3.19-3.55) in
TL. Dorsal fin low, its origin ahead of gill open-
ings, arising above fourth vertebra. Snout 6.37
(5.76; 5.29-6.37), upper jaw 3.01 (2.78; 2.65-3.01)
times in head length (HL). Eye 9.5 (9.4;
8. 3- 10.4) in HL and 1.5(1.63; 1.4-1.9) in snout,
closer to rictus than to tip of snout. Fleshy in-
terorbital width 7.8 (8.4; 7.7-9.9) in HL. Gill
openings nearly horizontal, their centers slightly
below midbody, their length about equal to di-
ameter of eye.

Anterior nostril tubular, elongate, slightly less
than eye diameter in length. Posterior nostril a
hole above eye, beginning in a line with eye.

Jaws subequal, the mouth closing completely.
Teeth in jaws uniserial, stout, pointed and slight-
ly retrorse. Six pairs of intermaxillary canines

form a U-shaped margin around three central
canines, the third the largest. Approximately six
uniserial, small vomerine teeth. About 12 upper
jaw teeth pairs, 18 lower jaw pairs; 3 pairs of
depressible canines behind mandibular sym-
physis.

Number of vertebrae 130 (129.7; 128-131),
50.5 (50.8; 50-51.5) before anal fin. First dorsal
pterygiophore arises above fourth vertebra.

Head pores present but not obvious. A single
pore anterior and proximal to, and a second pore
below base of anterior nostril. Six pores along
the mandible, the second through fifth the larg-
est. Four equally spaced pores along upper jaw,
the first beneath nostril base, the last beneath
rear of eye. A single pore between anterior and
posterior nostrils.

Color in isopropyl alcohol cream, overlain
with 17-21 distinctive brown bands which com-
pletely encircle head and body and extend onto

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 2

FIGURE 5. Uropterygius xanthopterus , CAS 35254, 245 mm TL, from Kapingamaringi, displaying white cephalic puncta-
tions. Arrows indicate the location of the anterior lateral line pores.

the Red Sea. Its synonyms also include Mu-
raena interrupta Kaup, 1856; Sideria chlevastes
Jordan and Gilbert, 1883; Gymnothorax signifer
Bliss, 1883; G. leucacme Jenkins, 1904; and G.
waialuae Snyder, 1904.

Whereas previous literature has suggested
that most morays are nocturnal, it now appears
that many, and possibly the majority of, moray
species are diurnal but rarely observed due to
their secretive nature (fide Chave and Randall
1971; Hobson 1974). On the basis of material
collected and our observations while diving, we
presume that G. enigmaticus is a shallow-water,
nocturnally active piscivore (Fig. 4). It is note-
worthy that G. rueppelliae is also a nocturnal
predator (Hobson 1974, as G. petelli), as is G.
undulatus (our observations), both of which are
also strongly banded species.

Uropterygius xanthopterus Bleeker, 1859

Uropterygius xanthopterus Bleeker, 1859, has
had a sketchy taxonomic history. We have lo-
cated the type-specimen, recognize it as a valid
species, and include U. alboguttatus Smith,
1962, in its synonymy.

Weber and de Beaufort (1916:397), without
comment, included U. xanthopterus in the syn-
onymy of Gymnomuraena marmorata La-
cepede, 1803, a wide-ranging, elongate Indo-Pa-
cific species of Uropterygius which possesses a
single anterior lateral line pore and lacks white
spotting on its head. Schultz (in Schultz et al.
1953:154) and Gosline (1958:226), on the basis

of central Pacific specimens, recognized U. xan-
thopterus as a distinct small species (the largest
of 2 1 3 specimens from 76 CAS rotenone collec-
tions in the Indian and central Pacific oceans
which we examined was 345 mm) which pos-
sesses two anterior lateral line pores and white
cephalic punctations (Fig. 5). Smith (1962:427)
again synonymized U. xanthopterus with U.
marmoratus and described U. alboguttatus on
the basis of Indian Ocean and Schultz' s central
Pacific specimens. In describing U. kamar
McCosker and Randall, 1977, we considered U.
alboguttatus to be a possible synonym of U.
xanthopterus. One of us (JEM) has subsequently
examined the complete type-series of U. albo-
guttatus and was unable to find differences in
coloration, meristic features, or morphometry.
The type-specimen of U. xanthopterus has
not been clearly identified; however, through
correspondence with Alwyne Wheeler, we have
located the 275-mm specimen in the British Mu-
seum (cat. no. 1867.11.28.271) received from
Bleeker and labeled "Muraena xanthopterus."''
In that no specimen similar to Bleeker' s type
exists in the Rijksmuseum (M. Boeseman,
in litt.), we presume that this is the type, and
the specimen which Bleeker illustrated and
described in his Atlas (1864:pl. CLXIV, fig.
4). A radiograph of the British Museum speci-
men clearly indicates that it is not U. marmo-
ratus, a species which possesses obvious, large
intramuscular bones.

McCOSKER & RANDALL: INDIAN OCEAN EELS

FIGURE 6. Uropterygius marmoratus, BPBM 12336, 701 mm, Nuku Hiva, Marquesas.

FAMILY OPHICHTHIDAE
Ophichthus retifer Fowler, 1935

Fowler (1935) described and illustrated Oph-
ichthus retifer on the basis of a 718-mm speci-
men from Durban, Natal. Eugenie Bohlke has
kindly examined the holotype (ANSP 63915) for
us and compared it with a syntype (ANSP 26224)
of O. erabo (Jordan and Snyder, 1901) from Ja-
pan. They do not significantly differ in color-
ation or proportions, yet there is a vertebral dif-
ference. A radiograph of the holotype of O.
retifer shows 143 vertebrae, with 73 before the
anal opening. McCosker (1979) reported that six
specimens of O. erabo from Japan, Hawaii, and
Taiwan had 152-155 vertebrae (x = 154). Fow-
ler (1935) suggested that O. retifer was "greatly
like Microdonophis fowleri Jordan and Ever-

mann 1903" (=O. erabo fide McCosker 1979)
"and its synonym Ophicthys garretti Giinther
1910" (a valid species). We consider O. retifer
to be conspecific with O. erabo, and account the
vertebral difference to clinal variation.

ACKNOWLEDGMENTS

We thank the following individuals: Susan
Middleton for photographic assistance; Michael
Hearne for the preparation of the radiographs;
M. Boeseman (Rijksmuseum van Natuurlijke
Historic), Eugenie and James Bohlke (ANSP),
William Fink (MCZ), W. Klausewitz (SMF),
Margaret M. Smith (RUSI), and Alwyne Wheel-
er (BMNH), for assistance with museum speci-
mens and records; Lillian Dempster and W. I.
Follett (CAS) for nomenclatural advice; and the

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 2

FIGURE 7. Adult Ophichthus erabo, from Jordan and Snyder (1901).

curators and staffs of many museums for allow-
ing us to examine specimens under their care.
Randall's collections were made possible in part
by grants from the National Geographic Society
and the American Philosophical Society. A por-
tion of McCosker's work was supported by
funds from the Charline Breeden Foundation.

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BLEEKER, P. 1859. Over eenige vischsoorten van de Zuid-
kustwateren van Java. Nat. Tijdschr. Neder. -Indie 19:329-
352.

. 1864. Atlas ichthyologique des Indes Orientales Neer-

landaises. Vol. 4. Amsterdam. 132 p.

BLISS, R. 1883. Descriptions of new species of Mauritian fish-
es. Trans. Soc. Roy. Arts Sci., Maurice 13:45-63.

BLOCH, M. E. 1795. Naturgeschichte der Auslandischen
Fische. Vol. 9.

CHAVE, E. H., AND H. A. RANDALL. 1971. Feeding behavior
of the moray eel, Gymnothorax pictus. Copeia 1971
(3):570-574.

FOWLER, H. W. 1935. South African fishes received from Mr.
H. W. Bell-Marley in 1935. Proc. Acad. Nat. Sci., Phila-
delphia 87:361^*08.

. 1956. Fishes of the Red Sea and southern Arabia. 1.

Branchiostomida to Polynemidae. Weizmann Sci. Press,
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GOSLINE, W. A. 1958. Central Pacific eels of the genus Urop-
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GUNTHER, A. 1910. Andrew Garrett's Fische der Siidsee,
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HOBSON, E. S. 1974. Feeding relationships of teleostean fish-
es on coral reefs in Kona, Hawaii. Fish Bull., U.S.
72(4):915-1031.

JENKINS, O. P. 1904. Report on collections of fishes made in
the Hawaiian Islands, with descriptions of new species.
U.S. Bur. Fish. Fish. Bull. 22:417-511.

JORDAN, D. S., AND C. H. GILBERT. 1883. Description of a

new muraenoid eel from the Galapagos Islands. Proc. U.S.

Natl. Mus. 6:208-210.
, AND J. O. SNYDER. 1901. A review of the apodal

fishes or eels of Japan, with descriptions of 19 new species.

Proc. U.S. Natl. Mus. 23:837-890.
KAUP, J. 1856. Uebersicht der Aale. Arch. Naturges.

22(l):41-77.
KLUNZINGER, C. B. 1871. Synopsis der Fische des Rothen

Meeres. II. Theil. Verh. Zool.-Bot. Ges. Wien 21:441-668.
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McCosKER, J. E. 1979. The snake eels (Pisces, Ophichthidae)

of the Hawaiian Islands, with the descriptions of two new

species. Proc. Calif. Acad. Sci., ser. 4, 42(2):57-67.
, AND J. E. RANDALL. 1977. Three new species of

Indo-Pacific moray eels (Pisces: Muraenidae). Proc. Calif.

Acad. Sci., ser. 4, 41(3): 161-168.
, AND R. H. ROSENBLATT. 1975. The moray eels

(Pisces: Muraenidae) of the Galapagos Islands, with new

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Bernice P. Bishop Mus. 24(11): 167-214.
RUPPELL, W. P. E. S. 1852. Verzeichniss der in dem Museum

der Senckenbergischen . . . Fische und deren Skelette.

Frankfurt-a-M.
SCHULTZ, L. P., AND COLLABORATORS. 1953. Fishes of the

Marshall and Marianas islands. Families from Asymme-

trontidae through Siganidae. U.S. Natl. Mus. Bull. 202, 1.

685 p.
SMITH, J. L. B. 1962. The moray eels of the western Indian

Ocean and the Red Sea. Ichthyol. Bull. Rhodes Univ.

23:421-444.
SNYDER, J. O. 1904. A catalogue of the shore fishes collected

by the steamer "Albatross" about the Hawaiian Islands in

1902. U.S. Bur. Fish. Fish. Bull. 22:513-538.
WEBER, M., AND L. F. DE BEAUFORT. 1916. The fishes of

the Indo-Australian Archipelago. Vol. 3. Leiden. 455 p.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 3, pp. 27-42

June 15, 1982

NEW SPECIES OF NORTH AMERICAN TACHYSPHEX WASPS
(HYMENOPTERA, SPHECIDAE)

By
Wojciech J. Pulawski

California Academy of Sciences, Golden Gate Park,
San Francisco, California 94118

ABSTRACT: The following new North American species of Tachysphex are described (type-localities are given
in parentheses): acanthophorus (Arizona: Willcox), apricus (California: Borrego Valley), arizonac (Arizona: 2
mi. NE Portal), armatus (Nevada: Sandy), bohartorum (California: Boca), idiotrichus (Arizona: 5 mi. W Portal),
irregularis (California: Hallelujah Junction), krombeiniellus (Florida: Levy County), lamellatus (Mexico: Sonora:
Alamos), menkei (California: Borrego Valley), mirandus (California: Palm Springs), musciventris (California:
Borrego), occidentals (California: 12 mi. E Lone Pine), papago (Arizona: Nogales), Solaris (California: Borrego
Valley), spatulifer (California: Arroyo Seco Camp), verticalis (California: 9 mi. W Beaumont), yuma (Mexico:
Baja California: La Paz), and vo/o (California: Davis).

INTRODUCTION

For several years, I have been working on a
monographic revision of North American
Tachysphex. Because of the size of this under-
taking, it will be some time before it is finished.
Therefore, I am describing some of the new
species now so their names will be available to
those persons working on Tachysphex behavior.
Furthermore, many hundreds of paratypes have
been deposited in 34 collections in the USA and
abroad, and it is desirable to validate these
manuscript names now to avoid their possible
use as nomina nuda in the works of others. The
descriptions given below are restricted to those
features which enable unambiguous recognition
of each species. More complete characteriza-
tions will be given when my revision is pub-
lished.

The terminology used below is based mainly
on Bohart and Menke (1976). A few terms which
need clarifications are the following:

clypeus: the clypeus has a midsection and two
lateral sections; the midsection usually has a
densely punctate, setose basomedian area, a
sparsely punctate shiny bevel, and a marginal
lip.

scutum: this term is used here for brevity's sake
instead of mesoscutum.

tergum, sternum: short terms for gastral tergum,
gastral sternum.

Many collectors are cited numerous times in
the lists of material examined. Their names have
been abbreviated to initials, as follows: ASM,
A. S. Menke; BV, B. Villegas; DRM, D. R.
Miller; EEC, E. E. Grissell; EIS, E. I. Schlin-
ger; GEB, G. E. Bohart; GDB, G. D. Butler;
FDP, F. D. Parker; FGW, F. G. Werner; FXW,
F. X. Williams; HKC, H. K. Court; JCH, J. C.
Hall; JAP, J. A. Powell; JMD, J. M. Davidson;
JWMS, J. W. MacSwain; LAS, L. A. Stange;
MAC, M. A. Cazier; MEI, M. E. Irwin; MSW,

[27]

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3

M. S. Wasbauer; PDH, P. D. Hurd; PMM, P.
M. Marsh; PFT, P. F. Torchio; RGB, R. C.
Bechtel; RMB, R. M. Bohart; ROS, R. O.
Schuster; RRD, R. R. Dreisbach; TG, Terry
Griswold; WJP, W. J. Pulawski.

In the geographic names below, the following
words have been abbreviated: County, Co.;
Creek, Cr.; Highway, Hwy.; Island, I.; miles,
mi.; Mountain(s), Mt(s).; River, R.; Station, Sta.
The name Lower California has been used for
the peninsula rather than Baja California, be-
cause the latter may refer either to the peninsula
or to a state in Mexico. Altitudes and distances
are given as they appear on the original labels —
in feet and miles. Multiplying the distances in
miles by 1.609 and the elevations in feet by
0.3048 will convert them into kilometers and
meters, respectively.

SOURCES OF MATERIAL AND

ACKNOWLEDGMENTS

The specimens described in this paper came
from institutional and private collections listed
below. The initials preceding the names are the
abbreviations by which institutions or private
collections are referred to in the text.

AMNH: American Museum of Natural History, New York,
New York (M. Favreau)

ANSP: Academy of Natural Sciences of Philadelphia, Phila-
delphia, Pennsylvania (D. C. Rentz)

ASU: Arizona State University, Department of Zoology,
Tempe, Arizona (F. F. Hasbrouck)

BMNH: British Museum (Natural History), London, England
(C. R. Vardy)

CAS: California Academy of Sciences, San Francisco, Cali-
fornia (P. H. Arnaud, Jr., T. J. Zavortink, W. J. Pulawski)

CIS: California Insect Survey, Division of Entomology, Uni-
versity of California, Berkeley, California (H. Daly)

CNC: Canadian National Collections of Insects, Arachnids
and Nematodes, Biosystematics Research Institute, Otta-
wa, Ontario (J. R. Barron)

CSDA: California Department of Food and Agriculture, Sac-
ramento, California (M. S. Wasbauer)

CSU: Colorado State University, Department of Zoology and
Entomology, Fort Collins, Colorado (H. E. Evans)

CU: Cornell University, Department of Entomology and Lim-
nology, Ithaca, New York (L. L. Pechuman)

FSCA: Florida State Collections of Arthropods, Gainesville,
Florida (E. E. Grissell)

HKT: H. K. Townes, American Entomological Institute, Ann
Arbor, Michigan

INHS: Illinois State Natural History Survey, Urbana, Illinois
(W. E. LaBerge)

KU: University of Kansas, Snow Entomological Museum,
Lawrence, Kansas (G. W. Byers)

KVK: K. V. Krombein, Arlington, Virginia (private collec-
tion), now in USNM

LACM: Natural History Museum of Los Angeles County, Los
Angeles, California (R. R. Snelling)

MCZ: Museum of Comparative Zoology at Harvard Univer-
sity, Cambridge, Massachusetts (J. Lawrence, J. C. Scott,
M. K. Thayer)

MPM: Milwaukee Public Museum, Milwaukee, Wisconsin (J.
K. Lawton)

NYSU: New York State University, College of Environmen-
tal Sciences and Forestry, Department of Environmental
and Forest Biology, Syracuse, New York (F. E. Kurczews-
ki)

OSDA: State of Oregon Department of Agriculture, Salem,
Oregon (R. L. Westcott)

OSU: Oregon State University, Department of Entomology,
Corvallis, Oregon (P. Oman, G. R. Ferguson)

TG: Terry Griswold, % Bee Biology & Systematics Labora-
tory, Utah State University, Logan, Utah (private collec-
tion)

UAE: University of Alberta, Department of Zoology, Ed-
monton, Alberta (A. L. Steiner)

UAT: University of Arizona, Department of Entomology,
Tucson, Arizona (F. G. Werner)

UCD: University of California, Davis, Department of Ento-
mology, Davis, California (R. M. Bohart, R. O. Schuster)

UCR: University of California, Riverside, Department of Bi-
ological Control, Riverside, California (S. Frommer)

UFG: University of Florida, Department of Entomology and
Nematology, Gainesville, Florida (B. Saffer)

UGA: University of Georgia, Department of Entomology,
Athens, Georgia (R. W. Matthews, C. L. Smith)

UIM: University of Idaho, Department of Entomology, Mos-
cow, Idaho (W. F. Barr)

UMSP: University of Minnesota, Department of Entomology
and Zoology, St. Paul, Minnesota (P. J. Clausen)

USNM: United States National Museum of Natural History
(Smithsonian Institution), Washington, D.C. (A. S. Menke,
K. V. Krombein)

USU: Utah State University, Department of Zoology, Logan,
Utah (G. E. Bohart, F. D. Parker, Terry Griswold)

WJP: Wojciech J. Pulawski, % California Academy of Sci-
ences, San Francisco, California (private collection)

WSU: Washington State University, Department of Entomol-
ogy, Pullman, Washington (M. T. James, R. Zack)

I express my sincere thanks to the curators
and other persons who kindly submitted speci-
mens for study. I feel especially indebted to R.
M. Bohart, A. S. Menke, K. V. Krombein, and
F. F. Kurczewski who helped me in many ways.

SPECIES GROUPS

Sixteen species groups are recognized in
Tachysphex (see Pulawski 1971, 1974, 1977), but
only four of them are represented in North
America. They are: the pompiliformis, termi-
natus, brullii, and julliani groups. The species
described in this paper belong to the pompili-
formis and brullii groups which are defined as
follows:

The pompiliformis group lacks peculiarities
which characterize other groups and thus pos-
sibly is a heterogenous assemblage of conve-
nience. The propodeal hindface in this group is
inclined, the female pygidial plate is not broad-

NEW SPECIES OF NORTH AMERICAN TACHYSPHEX

ened and without peculiar microsculpture, the
preapical bristles on the female gastral segments
are not thickened, and the male sterna are pru-
inose (except in mirandus). By comparison, in
the julliani group the propodeal hindface is ver-
tical or nearly so, male sterna are glabrous or
sparsely pruinose, and in the females of most
species the preapical bristles of gastral segments
IV and V are thickened, and the pygidial plate
is broadened or has a peculiar microsculpture.
The vertex is simple in the pompiliformis group,
while in the terminatus group a swelling is pres-
ent behind each hindocellus. Unlike the brullii
group, the apical female tarsomeres are simple
(see that group for details). The pompiliformis
group is cosmopolitan. Its species prey upon
acridid nymphs, but the Palearctic species ful-
vi tars is collects tettigonids. The following new
species are members of the pompiliformis
group: apricus, arizonac, bohartorum, idiotri-
chus, irregularis, lamellatus, mirandus, musci-
ventris, occidentalis, papago, Solaris, spatuli-
fer, verticalis, yolo, and yuma.

The brullii group is characterized by the pe-
culiar apical female tarsomeres: dorsum convex,
apicoventral margin produced into a lobe or at
least convex, and vertex variously modified
(covered with erect setae except glabrous ba-
sally, or angulate basally in lateral view, or
densely spinose). In other groups the dorsum is
scarcely convex, the apicoventral margin is
straight or nearly so, and the venter is evenly
covered with setae which are usually inclined
(but erect in verticalis), and it may have a few
spines in some species. Furthermore, the pro-
podeal dorsum setae are erect or inclined back-
wards in most species of the brullii group, but
only laterally so in acanthophorus, alayoi, ar-
matus, many individuals of mundus, and some
Australian species. Setae are inclined obliquely
cephalad in the Australian species brevicornis
and in most species of other groups. The brullii
group is widespread throughout all zoogeo-
graphic regions. Some species prey upon tetti-
gonids, while others are blattid collectors. The
following new species are members of this
group: acanthophorus, armatus, krombeiniel-
lus, and menkei.

SPECIES OF THE POMPILIFORMIS GROUP
Tachysphex apricus sp.n.

ETYMOLOGY. — The specific name apricus is
a Latin word meaning exposed to the sun.

DIAGNOSIS. — Tachysphex apricus differs from

other species of the pompiliformis group by the
setal pattern of its propodeal dorsum: median
setae are inclined cephalad, but the lateral setae
are directed obliquely backwards and join api-
comesally. Some species of the brullii group
(e.g., acanthophorus) have an identical pattern,
but the unspecialized apical female tarsomere of
apricus is distinctive. The male of apricus can
be distinguished by the compressed femoral
notch whose glabrous bottom forms an obtuse,
longitudinal crest. T. idiotrichus has a similar
crest, but unlike that species the body vestiture
is short in apricus. Unlike most species of the
pompiliformis group, the propodeal side of apri-
cus is alutaceous, shiny, impunctate or minutely
punctate.

GEOGRAPHIC DISTRIBUTION. — Xeric areas
between southern Texas, southern Nevada, and
southern California, and also Lower California.

MATERIAL EXAMINED. — HOLOTYPE: d, California, San
Diego Co., Borrego Valley, 3 May 1956, P. D. Kurd (UCD).

PARATYPES: 38 9 , 60 6 , 31 Mar. to 3 July, 10 and 31 Aug.,
9 Sep. Specimens for which institution is not indicated below
are all in UCD.

UNITED STATES OF AMERICA

Arizona. Cochise: 6 mi. N Apache, collector unknown (1
d, NYSU). Coconino: 4.5 mi. E Moenkopi, JMD & MAC (1
9, ASU). Maricopa: 10 mi. E Gila Bend, GDB (2 d); 3 mi.
sw Wickenburg, PFT & GEB (Id, USU). Mohave: 4 mi. w
Chloride, PFT, GEB, FDP (1 d, USU); 8 mi. E Mesquite
(Nevada), FDP & PFT (19, USU). Pfma: Organ Pipe Cactus
National Monument, J. L. Sperry (19,1 d); Tucson, W.
Benedict (19, NYSU), Bryant (1 9 , 1 d , CAS), FDP, LAS
(1 9, 2 d; 1 9, WJP). Final: w Stanfield, GDB & FGW (1
9). Yavapai: Bloody Basin, collector unknown (1 9); 10 mi.
NW Congress, FDP & LAS (19).

California. Imperial: Glamis, RMB (1 9), FDP(1 d); Palo
Verde, ROS (19); Pinto Flat, FXW (Id, CAS). Inyo: An-
telope Springs, HKC (19,5 d); Big Pine Cr., RMB (1 9),
FDP (1 d); 2 and 5 mi. E Big Pine, EEG (19,1 d); Little
Lake, BV (1 9); 3 mi. w Lone Pine, RMB (2 d); Tuttle Cr.
(2 mi. sw Lone Pine), JAP (1 9 , CIS). Kern: Kernville, T. R.
Haig (Id). Riverside: 18 mi. w Blythe, RMB (Id); 3.5 and
4 mi. s Palm Desert, MEI, S. Frommer & R. M. Worley (2
9 , UCR); San Andreas Canyon, RMB (Id); Shavers Summit,
MSW (1 9 , UCD); San Timoteo Canyon, MSW, R. McMaster
(1 9, CSDA). San Bernardino: 1 mi. s Adelanto, MEI (1 d);
Colton Hills, TG (2 9 , TG), Kramer, MSW (29,3d, CSDA);
3 mi. s Kramer Junction, MEI (1 9); sand dunes 7 mi. sw
Kelso, MSW & J. S. Wasbauer (1 9 , CSDA), Mitchells Cav-
erns, TG (1 9 , TG), 36 road mi. E Twenty nine Palms, TG (1
d , TG). San Diego: Borrego Valley, RMB (4 d ; 1 9 , 2 d ,
USNM), JCH (Id), PDH (4 d), G. A. Marsh (19), EIS (2
d; 2 d, WJP), MSW (Id, CIS), MSW, J. Slansky (19,1
d , CSDA), FXW (9 d , CAS); Scissors Crossing, J. C. Down-
ey (1 9), H. & M. Townes (Id, HKT). Ventura: Sespe Can-
yon, R. W. Sporne (Id).

Nevada. Clark: Jean, GEB (3 d); 30 mi. s Searchlight, PFT,
R. Rust, Youssef (1 d, USU).

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3

New Mexico. Dona Ana: Las Cruces, RMB (19, UCD).
Otero: Alamogordo, collector unknown (19); Alamo Canyon
near Alamogordo, MEI (Id, UCR).

Texas. Brewster: Big Bend National Park (Nine Point
Draw), W. R. M. Mason (19, CNC).

MEXICO

Lower California. San Angel, N. Leppla, JMD, J. Bigelow,
M. Bentzien, W. Fox, S. William, MAC (19, ASU); 16 mi.
N Puertocitos, MEI (19, UCR).

Tachysphex arizonac sp.n.

ETYMOLOGY. — The specific name arizonac is
an Indian word meaning little spring.

DIAGNOSIS. — Tachysphex arizonac is charac-
terized by the well-defined mesopleural punc-
tures, median scutal setae transversely oriented,
and sternum I with a horizontal depression api-
cally. The females of arizonac and lamellatus
have a peculiar clypeus whose free margin is
undulate; they can scarcely be distinguished
from each other. A useful character is the hy-
postomal carina which is low in arizonac, but
high in many lamellatus. Furthermore, in some
arizonac the middle projection of the clypeal lip
is markedly larger than the sublateral one (pro-
jections about equal in lamellatus). The male of
arizonac has a peculiar clypeus: bevel semilu-
nate, lobe forecorner prominent (acutely in
some specimens), lip usually with obtuse pro-
jection mesally. The clypeus is somewhat simi-
lar in texanus, but in that species setae are ap-
pressed or nearly so beneath the mesopleural
scrobe (setae suberect in arizonac).

GEOGRAPHIC DISTRIBUTION. — Southern Utah,
Arizona, and adjacent areas of California; So-
nora State in Mexico.

MATERIAL EXAMINED. — HOLOTYPE: <S , Arizona, Cochise
Co., 2 mi. NE Portal, 26 June 1964, J. M. Puckle, M. A. Mor-
tenson, M. A. Cazier (UCD).

PARATYPES: 13 9 , 17 d , 5 May to 17 July.

UNITED STATES OF AMERICA

Arizona. Cochise: Huachuca Mts., FXW (1 9, CAS; 1 9,
3 6 , UCD). Maricopa: 3 mi. sw Wickenburg, PFT & GEB
(19, UCD). Pima: Brown Canyon, Baboquivari Mts., K. W.
Radford & W. Patterson (1 9, 2 d, UAT); Sabino Canyon,
Santa Catalina Mts., RMB (1 <S , UCD), collector unknown
(19, WJP).

California. Imperial: Experimental Farm and no specific lo-
cality, J. C. Bridwell (3 9 , 1 d , USNM; 2 9 , 1 d , WJP).
Inyo: Big Pine, N. J. Smith (1 6", UCD); Deep Springs, H.
Nakakihara (1 3 , UCR). Riverside: Upper Deep Canyon at
Horsethief Cr., MEI (1 d, UCR).

Utah. Washington: Leeds Canyon, G. F. Knowlton, W. J.
Hanson, T. H. Hsiao (2 9, 2 3, USU; 1 9, WJP); Santa

Clara, FDP & PFT (Id, USU); Toquerville, G. F. Knowlton,
W. J. Hanson (1 d, USU).

MEXICO

Sonora. Cocorit, FDP & LAS (Id, UCD); La Aduana,
FDP& LAS(1 d, UCD).

Tachysphex bohartorum sp.n.

ETYMOLOGY. — This species is dedicated to
Dr. G. E. Bohart (Logan, Utah) and Dr. R. M.
Bohart (Davis, California) who have made ex-
tensive contributions to our knowledge of North
American Tachysphex, and who have also
helped me in my studies.

DIAGNOSIS. — Like pechumani, the female of
bohartorum has brassy golden frontal vestiture.
The frontal vestiture is also golden in psilocerus
which has a peculiar upper metapleuron. Unlike
pechumani, the gaster of female bohartorum is
all red, the flagellum is all black, the clypeal lip
is deeply indented laterally (shallowly in pechu-
mani), and the mesothoracic venter is peculiar
(the posterior, horizontal part is shorter than the
anterior, oblique part); the densely punctate-
throughout tergum V is a subsidiary recognition
feature. T. bohartorum is known from Califor-
nia, Oregon, and Nevada, while pechumani oc-
curs in New Jersey and Michigan.

The male of bohartorum can be recognized by
the closely punctate terga (punctures mainly
subcontiguous) combined with the apicomesally
impunctate and glabrous sterna II-IV.

Supplementary diagnostic characters of both
sexes are: propodeum not ridged (at most the
hindface has a few, inconspicuous ridges be-
low); propodeal dorsum with setae appressed
mesally, oriented obliquely anterad.

GEOGRAPHIC DISTRIBUTION. — Montane areas
of southern Oregon and northern California, also
Sierra Nevada.

MATERIAL EXAMINED. — HOLOTYPE: 9 , California, Nevada
Co., Boca, 11 July 1961, R. M. Bohart (UCD).

PARATYPES: 90 9 , 23 1 d , 24 Apr. , 8 June to 14 Aug. , mainly
UCD, also BMNH, CAS, CIS, CNC, CSDA, LACM, MCZ,
OSU, TG, UCR, USNM, WJP.

California. Alpine: Carson Pass, RMB (Id); Hope Valley,
PDH (1 d), P. C. Hutchinson (1 9); 15 mi. NE Red Lake, EIS
(2 d); Winnemucca Lake, RMB (19,2 d), N. J. Smith (2
9, 1 d); Woodfords, W. W. Middlekauf (3 d). Del Norte:
Little Grayback (NE part of county), JAP (4 9 , 4 d , CIS).
Eldorado: Meyers, RMB (1 d); Tahoe, FXW (39,3 d);
Strawberry Valley, E. C. VanDyke (1 9); Echo Lake, W. W.
Middlekauf (1 9 , 1 d); Echo Portal, P. H. Arnaud (1 9); Echo
Pass, C. A. Downing (19). Humboldt: Red Cape Lake, TG

NEW SPECIES OF NORTH AMERICAN TACHYSPHEX

(1 9, 1 d, TG). Inyo: Big Pine Cr., 7500 ft, RMB (19,1
d); near Mono Pass, 12,000 ft, C. D. MacNeill (1 9, CIS).
Lassen: Bridge Cr. Camp, RCB, RMB, G. Schaefers (69,1
<J); Summit Camp, JWMS (1 9). Mariposa: Sentinel Dome
Cutoff (Yosemite National Park), T. N. Seeno (19). Modoc:
Cedar Pass in Warner Mts., JWMS (1 9 , 2 d , CIS), 6000 ft,
collector unknown (19); Warner Mts. 2 mi. NNW Fort Bid-
well, D. C. Rentz & C. D. MacNeill (1 d, CAS). Mono: 11 mi.
N Bridgeport, RMB, PMM (3 d); East Walker R., 13 mi. NE
Bridgeport, MSW & J. S. Wasbauer (2 d); Leavitt Meadow,
R. L. Usinger (1 6. CIS). Nevada: Boca, RMB, MEI, PMM,
WJP (89, II d); Sagehen Cr. near Hobart Mills, RMB, R.
C. Blaylock, R. L. Brumley, M. A. Chambers, R. H. Good-
win, EEG, D. S. Horning, PDH, MEI, JAP, WJP, G. Schae-
fers, R. L. Westcott (15 9 , 98 d). Placer: Carnelian Bay (Lake
Tahoe), RMB, FDP, WJP, BV (3 9, 21 d); Tahoe, FXW (1
d). Plumas: Bucks Lake, EIS (1 d); Burks, FXW (1 9); Lake
Almanor, E. G. Linsley (1 d); Meadow Valley, W. H. Nelson
(1 9); 14 mi. w Quincy, W. Turner (1 9 , 2 d , WSU). Shasta:
Lake Eiler, C. H. Spitzer (1 9); Lassen Peak, 7500ft, JWMS
(1 9). Sierra: Independence Lake, RMB, R. D. Moon (16
9 , 51 d), BV (1 9); Kyburz Flat, RMB (Id); Sierra Buttes,
F. J. Montgomery (19); Sierra Valley, RMB (Id); Sierra-
ville, RMB (1 d); Yuba Pass, MEI (1 d, UCR). Siskiyou:
McBridge Springs (3 mi. NNE Mt. Shasta City), C. D. MacNeill
(3 d); Mount Shasta, JAP (2 9), 1 mi. SE Salmon Mt., TG (2
d, TG). Trinity: Coffee Cr. Ranger Sta., A. J. Mueller (1
9), JWMS (1 9, CIS). Tuolumne: Chipmunk Flat, ASM (1
9), JWMS (19, CIS); Dardanelle, EIS (19); Sonora Pass,
9-10,000 ft, S. M. Kappos, JWMS (1 9, 2 d).

Nevada. Douglas: Spooners Lake N junction Hwy. 28, P.
Adams (2 d). Washoe: Mount Rose, 6500 ft, RMB (Id).

Oregon. Jackson: 8 mi. SE Butte Falls, R. L. Westcott (1
9 , OSDA). Klamath: 15 mi. NE Ely, Schuh & Denning (19,
OSU); Eagle Ridge near Klamath Lake, C. L. Fox (1 9); Lake
of the Woods, H. A. Scullen (2 9). Lake: Warner Pass, DRM
d d).

Tachysphex idiotrichus sp.n.

ETYMOLOGY. — The specific name idiotrichus
is derived from the Greek words idios, own, pe-
culiar, and trix (tricho-), hair; with reference to
the peculiar vestiture.

DIAGNOSIS. — Tachysphex idiotrichus differs
from other North American species of the pom-
piliformis group by the unusually long setae on
the head, thorax, and femora (the vertex setae,
for example, equal 2.2-3.0 midocellar diame-
ters); the large punctures on the middle section
of the female clypeus (also basally); the pres-
ence of graduli on sterna III-V of the female;
and sparsely punctate (except apex) male ter-
gum VII. Subsidiary diagnostic characters are:
sparsely punctate vertex; and in the male: vertex
width more than twice length (like pechumani);
presence of graduli on sterna III-VI (like ash-
meadii, glabrior, irregularis, and verticalis); and
compressed forefemoral notch (as in apricus).

GEOGRAPHIC DISTRIBUTION. — Low moun-
tains of southwestern Texas, New Mexico,
southern Arizona, and southern California south
to Jalisco State, Mexico.

MATERIAL EXAMINED. — HOLOTYPE: 9 , Arizona, Cochise
Co., Southwest Research Sta., 5 mi. w Portal, 20 Oct. 1962,
Vincent Roth (UCD).

PARATYPES: 25 9 , 1 1 d ; 19 Feb. to 30 Nov.

UNITED STATES OF AMERICA

Arizona. Cochise: 3 mi. E Apache, Rozen & Schrammel (1
9, UCD); 2 mi. w Chiricahua National Monument, A. L.
Steiner (1 9, WJP); NW Portal, O. W. Richards (2 9,
BMNH); 2 mi. sw Portal, A. L. Steiner (1 9, UAE); same
data as holotype, except 1 9, 19 Feb. 1963 (3 9; UCD,
USNM, WJP); same locality, MAC & Ordway (1 d AMNH);
Sulphur Springs Valley, G. Munson (1 9, UCD); 14 mi. w
Tombstone, RMB (19, AMNH; 4 9 , 2 d , UCD; 1 d , USNM;
1 d, WJP); same locality, FDP (1 9, 2 d, UCD); 1 mi. SE
Willcox, G. I. Stage (19, CAS). Puna: Tucson, M. L. Noller
(19, UAT). Yavapai: Cottonwood, R. C. Miller (2 9 , 1 d ,
UCD); 7 mi. N Cottonwood, R. C. Miller (1 d, UCD).

California. San Bernardino: Mid Hills (9 mi. SSE Cima),
5400-5500 ft, TG (2 9 , TG).

New Mexico. Hidalgo: Rodeo to Road Forks (19, UCD).
Otero: Alamogordo, collector unknown (2 9 , UCD). Socorro:
10 mi. w Socorro, U. N. Lanham (1 9 , CSU).

Texas. Pecos: no specific locality, RMB (Id, UCD).

MEXICO

Jalisco. Lagos de Moreno, R. C. Bechtel & EIS (1 d,
UCD).

Tachysphex irregularis sp.n.

ETYMOLOGY. — The specific name irregularis
is a Latin word for irregular, with reference to
the mesopleural sculpture.

DIAGNOSIS. — Tachysphex irregularis differs
from other members of the pompiliformis group
in having a rugose or punctatorugose mesopleu-
ron. The subsidiary recognition features are: the
almost impunctate venter of the female trochan-
ters, and in the male: presence of graduli on ster-
na III-VI, sternal punctures about as large as
those on the mesothoracic venter.

GEOGRAPHIC DISTRIBUTION. — Western U.S.,
eastwards to Wyoming and New Mexico.

MATERIAL EXAMINED. — HOLOTYPE: 9 , California, Lassen
Co., Hallelujah Junction, 12 July 1954, G. A. Schaefers
(UCD).

PARATYPES: 41 9 , 29 d ; 16 Apr. to 1 1 Sep.

Arizona. Cochise: 8 mi. NE Apache, PDH, E. G. Linsley
( 1 2 , 1 d , CIS); 14 mi. w Tombstone, FDP (Id, UCD). Pinal:
8 mi. SE Olberg, M. A. Mortenson, JMD, MAC (12, UCD).

California. Inyo: Bishop, F. P. VanDuzee (2 9, UCD,
WJP). Lassen: Hallelujah Junction, RMB, J. E. Gillaspy, C.
J. Horning, G. A. Schaefers, F. Morishita, EIS, R. H. James,

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3

R. L. Sisson (11 9, 10 d; AMNH, MCZ, UCD, USNM,
WJP). Modoc: 5.5 mi. E Cedarville, V. L. Vesterby (Id,
UCD); Hot Cr., RMB & GEB (1 9, UCD). Mono: Benton
Crossing, F. G. Andrews (2 9 , CSDA). Nevada: Boca, RMB
(5 9, 1 d, UCD), MEI (Id, UCD); Sagehen Cr., PDH (I
d, UCD). Sierra: Sattley, RMB (2 d, UCD). Siskiyou: Mac-
doel, J. Schuh (2 9, UCD), Hatfield, J. Schuh (Id, UCD);
Mt. Shasta City, JAP (19, UCD); Red Rock, J. Schuh (1
9 , UCD). Tuolumne: Sonora Pass, R. W. Thorp (19, UCD).

Colorado. Jackson: 10 mi. N junction of roads 14 & 40, R.
& K. Dreisbach (19, UCD). Mineral: South Clear Cr., TG
(19, TG). Routt: 7 mi. E Hayden, J. S. Buckett (1 d , UCD).

Idaho. Canyon: Nampa, GEB (19, UCD). Cassia: 4 mi. SE
Malta, R. L. Westcott (1 9, UIM); 5 mi. N Malta, R. A.
Mackie (19, UIM). Custer: 2 mi. E Leslie, D. S. Horning (1
9 , UCD). Elmore: 4 mi. E Orchard, A. J. Walz (Id, UIM).
Owyhee: Silver City, A. R. Gittins (1 9 , UIM); 17 mi. w Silver
City, A. R. Gittins ( 1 9 , 1 d , UIM).

Nevada. Elko: Cobb Cr., 6 mi. sw Mountain City, P. H.
Baldwin (19, UCD). Humhol.lt: Orovada, MEI (Id, UCD),
FDP (19, UCD); Paradise City, P. H. Baldwin (Id, UCD);
15 mi. E Paradise City, collector unknown (Id, WJP). Wash-
oe: Reno Hot Springs, C. N. Slobodchikoff (1 9, CAS), Sky
Ranch near Reno, EIS (1 d, UCD).

New Mexico. Dona Ana: Las Cruces, Arabella Sanchez (1
9, UCD). Hidalgo: Rodeo, PDH (1 d, UCD), H. V. Weems
(Id, FSCA).

Oregon. Klamath: Lower Klamath Lake, J. Schuh (1 9,
UCD).

Wyoming. Fremont: Shoshoni. GEB & PFT (1 d, USU).

Tachysphex hum-Hat us sp.n.

ETYMOLOGY. — The specific name lamellatus
is a Latin adjective derived from lamella, with
reference to the unusually high hypostomal ca-
rina.

DIAGNOSIS. — Tachysphex lamellatus has a
punctate mesopleuron, median scutal setae
transversely oriented, and sternum I apically
with a horizontal depression. Several other
species share these features. Males of lamella-
tus and sonorensis have a triangular or subtrian-
gular clypeal lip and a nondentate inner mandib-
ular margin; unlike sonorensis, flagellomeres III
and IV of lamellatus are about equal in length.
Females of lamellatus and arizonac have a pe-
culiar, undulate-free margin of the clypeal lip,
but they cannot be distinguished with certainty
from each other. The hypostomal carina is un-
usually high in some females and most males of
lamellatus, and the gena adjacent to the hypo-
stoma is ridged in most males. Both features are
unique to lamellatus, and when present, distin-
guish the species from all other Tachysphex.

GEOGRAPHIC DISTRIBUTION. — New Mexico
to southern California, north to southwestern
Utah, and Mexico.

MATERIAL EXAMINED. — HOLOTYPE: d, Mexico, Sonora,
Alamos, 25 Feb. 1963, P. H. Arnaud, Jr. (CAS Type No.
13465).

PARATYPES: 76 9 , 45 d , 2 Jan., 25 Feb. to 15 May (Mexico),
10 Mar. to 25 Oct. (USA). Specimens for which institution is
not indicated below are kept in UCD.

UNITED STATES OF AMERICA

Arizona. Cochise: Paradise, LAS (1 9); Portal, JMD (1
9); 5 mi. sw Portal, C. W. Sabrosky (19, USNM); same
locality, M. Stathem (1 9); 5 mi. w Portal, V. Roth (1 d,
NYSU); same locality, collector unknown (1 d, NYSU);
Ramsey Canyon, Huachuca Mts., R. F. Sternitzky (3 d,
CNC). Gila: Gila R. 3 mi. sw Christmas, FGW (Id, UAT).
Pima: Baboquivari Mts., Bryant (19, CAS), A. L. Melander
(29,1 d), K. W. Radford & W. Patterson (7 9 , 2 d , UAT);
Elkhorn Ranch, Baboquivari Mts., M. L. Noller, K. Roever
(2 9 , UAT); Madera Canyon, 4400 ft, Santa Rita Mts., HKC
(1 9), ASM & WJP (1 9, WJP), FGW (3 9, 1 d, UAT);
Madrona Ranger Sta., Rincon Mts., M. L. Noller, J. C. Be-
quaert, H. Elton (2 9 , 1 d , UAT); Molino Camp, Santa Cat-
alina Mts., R. S. Beal (Id, CIS); Sabino Canyon, Santa Cat-
alina Mts., RMB (1 9), R. H. & L. D. Beamer, W. LaBerge,
C. Liang (19, KU), A. D. Telford (1 9 , 1 d); Santa Catalina
Mts.,FGW&GDB(l 9); Tucson, Bryant (1 9 , CAS), FGW,
Malaise trap (2 9 , UAT); 5 mi. N Tucson, FGW & GDB (1
9). Final: Superior, Boyce Thompson Arboretum, FGW, J.
Bequaert (Id, UAT). Santa Cruz: Florida Canyon (Santa Rita
Mts.), D. P. Levin (19, UIM); Madera Canyon (Santa Rita
Mts.), D. P. Levin (Id, UIM); Patagonia, FGW & GDB (1
9); Sycamore Canyon, Tumacacori Mts., A. & H. Dietrich
(19, NYSU). Yavapai: Irving Power Sta., w Strawberry, O.
Flint & ASM (Id, USNM). Also Atascasa Mts. [a mistake
for Atasco Mts.?], R. H. Crandall (1 9, 1 d, USNM).

California. Inyo: Darwin Falls, JAP (Id, CIS). Orange:
Laguna Canyon, MEI (1 9, UCR). San Bernardino: Cajon
Pass, J. E. Gillaspy (19); Mid Hills (9 mi. SSE Cima), TG (2
9 , 2 d , TG).

New Mexico. Catron: Glenwood, RMB (19). Luna: 6 mi.
NW Florida, J. S. Buckett, M. R. & R. C. Gardner (2 9).

Utah. Washington: Leeds Canyon, Malaise trap (29,1
d, USU).

MEXICO

Chihuahua. 6 mi. s Encinillas, MEI (19, UCR).

Hidalgo. Jackala, L. D. Beamer (1 9 , KU).

Puebla. 3 mi. NW Petlalcingo, FDP (I d).

Sinaloa. 9 mi. E Chupaderos, FDP, LAS (1 9 , 2 d); 54 mi.
s Culiacan, MEI (7 9 , 2 d , UCR).

Sonora. Alamos, same data as holotype (99, 17 d; CAS,
UCD, WJP); La Aduana, FDP, LAS (8 9 , 1 d ; 4 9 , UCR),
LAS (4 d).

Tres Marias Islands. Maria Magdalena L, H. H. Keifer (1
9 , CAS).

Tachysphex mirandus sp.n.

ETYMOLOGY. — The specific name mirandus
is a Latin word meaning wonderful, strange, sin-
gular.

DIAGNOSIS. — Tachysphex mirandus can be
recognized by its peculiar sculpture. In both

NEW SPECIES OF NORTH AMERICAN TACHYSPHEX

sexes, the punctures are larger on the lateral
clypeal section than on the adjacent frons, al-
though the difference is slight in some speci-
mens. The midfemur (also female forefemur) is
alutaceous basoventrally, with only a few,
sparse punctures. In the female, terga II-V are
alutaceous, glabrous (except somewhat pubes-
cent laterally). The male can also be recognized
by the mesally nonpubescent, largely glossy and
sparsely punctate sterna II-VI combined with
the nondentate inner mandibular margin, trian-
gular clypeal lip, and the nonangular clypeal free
margin between the lip and the lateral section.
Subsidiary recognition features of both sexes
are: mesopleural setae suberect (almost as in
semirufus); horizontal part of mesothoracic ven-
ter in most specimens glossy, sparsely punctate,
contrasting with dull, strongly microsculptured
mesopleuron.

GEOGRAPHIC DISTRIBUTION. — Xeric areas of
Nevada, Arizona, southern California, and
Lower California.

MATERIAL EXAMINED. — HOLOTYPE: 9 , California, San
Bernardino Co., Palm Springs, 11 Feb. 1958, A. Melander
(USNM).

PARATYPES: 17 9, 6 d, 20 Jan. to 14 Apr.

UNITED STATES OF AMERICA

Arizona. Mohave: 4 mi. s Hoover Dam, A. R. Gittins (1
d, UIM).

California. Fresno: Pinoche, 29 Mar. 1930, E. C. Van-
Dyke (1 $ , UCD). Imperial: 9 mi. w Coyote Wells, 26 Mar.
1961, PDH (Id, CIS); Ocotillo, 22 Mar. 1966, PDH (29,1
d, CIS); Yuha Desert, 15 Feb. 1948, C. D. MacNeill (1 9,
CAS). Kern: 3 mi. NW Indian Wells, 12 Apr. 1954, J. M.
Linsley (1 9, CIS); Short Canyon (7 mi. NW Inyokern), 15
Mar. 1955, PDH (Id, CIS). Los Angeles: Little Rock, 28 Mar.
1971, RMB (1 9, UCD), 22 Mar. 1941, Don Wasem (1 9,
UCD). Riverside: Whitewater, 14 Apr. 1958, H. R. Moffitt (1
9 , UCD). San Bernardino: 5 mi. s Essex, 26 Mar. 1970, EEG
& R. F. Denno (1 9 , UCD); 14 mi. E Newberry, A. E. Mich-
elbacher, 31 Mar. 1964 (1 9 , CIS). San Diego: Borrego Valley,
2 & 11 Apr., RMB (2 9; UCD, WJP); Borrego Springs, 30
Mar. 1960, MSW (Id, UCD). San Luis Obispo: Cuyama Val-
ley, 30 mi. w Maricopa, 2i Mar. 1931, E. P. VanDuzee (1
9 , UCD). Tulare: Kaweah Power House, 20 Jan. 1972, F. G.
Andrews (1 9, CSDA).

Nevada. Nye: Mercury, N. T. S. [collector ?], 24 Apr. 1961
(19, USNM), 28 Mar. 1962 (Id, USNM).

MEXICO

Baja California Norte. Diablo Canyon, E face of Sierra San
Pedro Martfr. 6 Apr. 1973, J. Donohoe (19, CIS); Upper
Camillas Canyon (Sierra Juarez), 19 Mar. 1967, JAP (19,
CIS).

Tachysphex musciventris sp.n.

ETYMOLOGY. — The specific name musciven-
tris is derived from the Latin words muscus,
moss, and venter; with reference to the dense
pubescence on the female mesothoracic venter
and male sterna.

DIAGNOSIS. — The peculiar mesothoracic ven-
ter of the female of musciventris is unique
among Tachyspex: the mesothoracic venter is
deeply sunken and densely pubescent along the
midline on the posterior (horizontal) half. The
mesally notched clypeal lip and the usually red
hindleg are additional diagnostic characters.

The males of musciventris and an undescribed
species have a triangular clypeal lip, ill-defined
mesopleural punctures, and velvety pubescent
sterna III-VI (integument all or largely con-
cealed). Unlike the other species, the vertex
width in musciventris is 1.6-2.0 times its length,
the vestiture is appressed on the hypoepimeral
area, and sternum II is velvety pubescent.

GEOGRAPHIC DISTRIBUTION. — California to
southwestern Texas, north to southwestern
Utah, south to northwestern Mexico.

MATERIAL EXAMINED. — HOLOTYPE: 9 , California, San
Diego Co., Borrego, 2 Apr. 1973, R. M. Bohart (UCD).

PARATYPES: 225 9, 216 d; 25 Mar. to 15 June, and 16-22
Nov.; mostly UCD, but also AMNH, BMNH, CAS, CIS,
CNC, CSDA, OSU, TG, UCR, UIM, USNM, USU, WJP.

UNITED STATES OF AMERICA

Arizona. Cochise: Dragoon Mts., GDB & FGW (19). Gila:
Globe, GEB (19). Maricopa: 8 mi. s Buckeye, MEI (3 d,
UCR); Gila Bend, GDB & FGW ( 1 9 ); 18 mi. s Gila Bend, S.
A. Gorodenski, JMD, MAC (1 9); 5 mi. N Mesa, A. R. Gittins
(19, UIM). Mohave: 8 mi. E Mesquite, Nevada, FDP & PFT
(3 9); 16 mi. N Wikieup, GDB & FGW (1 d). Final: w Stan-
field, GDB & FGW (Id). Santa Cruz: Tubac, PFT (Id).
Yavapai: 18 mi. N Aguila, FDP (1 9 , USU). Yuma: Dateland,
GDB & FGW (19); Ligurta, RMB (1 9); 15 mi. N Yuma,
MEI, FDP (2 9); 22 mi. N Yuma, S. A. Gorodenski, JMD,
MAC(1 9).

California. Amador: 5 mi. E Jackson, W. E. Simonds (1
d). Fresno: 10 mi. w Coalinga, RMB (1 d). Imperial: Choc-
olate Mts., Ogilby Road, 3 mi. s junction Hwy. 78, MSW (1
9, CSDA); Fish Cr. Mts., D. F. Hardwick (19); Glamis,
RMB, PMM, FDP (4 9 , 5 d); 3 mi. N Glamis, MJW (5 9);
Palo Verde, MSW & J. S. Wasbauer, PDH (3 9); 3 mi. s Palo
Verde, G. Tamaki (1 9 , CIS); 8 mi. s Palo Verde, C. A. Toschi
(Id, CIS). Inyo: Wildrose Canyon, ASM (Id). Lake: Lu-
cerne, D. J. & J. N. Knull (19). Kern: 14 mi. N Blackwells
Cor., C. D. MacNeill (2 d , CIS); Iron Canyon (El Paso Mts.),
C. A. Toschi (1 9, CIS); 19 mi. wShafter, RMB (1 9). Lassen:
Bridge Cr. Camp, J. E. Gillaspy (Id); Summit Camp, PDH,
E. G. Linsley (4 d). Los Angeles: 2.5 mi. Pearblossom, R. W.
Brooks (19); Little Rock, E. P. VanDuzee (19). Modoc:

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3

Cedar Pass, D. L. Dahlsten (16). Monterey: Monterey, FDP
(1 9). Riverside: Boyd Desert Research Center (4 mi. s Palm
Desert), C. A. Toschi (2 9 , CIS); 18 mi. w Blythe, RMB, D.
S. Horning, MEI, FDP, ROS (11 9 , 28 d); 10 mi. NW Cot-
tonwood (Joshua Tree National Monument), PDH (19, CIS);
Deep Canyon (3.5 mi. s Palm Desert), JCH (19, UCR), C.
Wilkinson (1 9); 5 mi. s Hemet, RMB (1 <S); Hwy.
74 x Strawberry Cr., EIS (1 9, CIS); Hopkins Well, PDH,
E. G. Linsley, JAP (6 9 , 1 1 d); Indio, J. Wilcox (19, OSU);
Keen Camp in San Jacinto Mts., EIS (1 d); Millard Canyon,
MEI (1 d); 7 mi. w North Palm Springs, TG (1 d, TG); Palm
Canyon, EIS (1 d, UCR); Palm Springs, C. Dammers, 11
Nov. (4 9); Pinon Flat in San Jacinto Mts., E. S. Ross (1
9); Riverside, JCH (19, UCR), TG (1 d, TG); 4 mi. s Riv-
erside, RCB, EIS (29,3 d); Shavers Summit, MSW (2 9);
Strawberry Canyon, W. A. Hunt (1 9, UCR); Thousand
Palms, RMB, EEG, MEI, PMM, DRM, FDP, W. R. Richards
(22 9, 51 d); 10 mi. E Whitewater RMB (19,3 d). San
Bernardino: 13 mi. E Amboy, E. G. Linsley, JWMS (13 9,7
d); Baker, F. G. Andrews (19, UCR); 4 mi. s Baker, MSW
& J. S. Wasbauer (1 9, 1 d); 3 mi. w Barstow, Brown &
Lundgren (1 9 , CAS); 2 mi. w Cajon Pass, EIS (1 d); Colton,
E. P. VanDuzee (1 9 ); 5 mi. SE Hesperia, Rozen & Schrammel
(2 9 ); sand dunes 7 mi. SE Kelso, MSW (1 d); Needles, JWMS
(1 d); 2 mi. w Phelan, EIS (1 d); Red Mountain, D. F. Hard-
wick (19); Vidal Junction, PDH (19). San Diego: Alpine,
FXW (1 d, CAS); Borrego, RMB, C. Goodpasture, EEG,
PDH, H. L. McKenzie, A. L. Melander, FDP, R. Snelling,
MSW (97 9, 56 d); Coyote Cr. (Borrego Valley), FDP (1
d); Coronado, F. E. Blaisdell (19); Scissors Crossing, R. R.
Pinger (1 d). San Luis Obispo: 5 mi. w Nipomo, RMB, C.
Goodpasture (5 d); 10 mi. w Simmler, PDH, R. W. Thorp,
C. A. Toschi (4 9 , 6 d , CIS). Ventura: Hungry Valley (5 mi.
s Gorman), PDH, JAP (7 9 , 7 d).

Nevada. Clark: 20 mi. w Glendale, FDP & PFT (4 9).

New Mexico. Dona Ana: 4 & 1 1 mi. N Las Cruces, 9 . W.
Richards (2 9); Mesilla, GEB (Id, UCD).

Texas. Hudspeth: Sierra Blanca, J. O. Martin (1 9).

Utah. Washington: Santa Clara, FDP & PFT (19, USU).

MEXICO

Lower California. 7 mi. s Guadalupe, MEI (Id, UCR), San
Quintin, FXW(1 9, CAS).

Sonora. 60 mi. E San Luis, G. R. Ballmer (1 9, UCR), 23
km sw Sonoita, B. & C. Durden (19, AMNH).

Tachysphex occidentalis sp.n.

ETYMOLOGY. — The specific name occidenta-
lis is a Latin adjective meaning western; with
reference to the geographic distribution of the
species.

DIAGNOSIS. — Sternal punctures of male occi-
dentalis are well defined, about as large as those
of the mesothoracic venter (sometimes mark-
edly larger). Sternal punctures are similar in
some other species (e.g., irregularis, tarsatus),
but the clypeal lobe of occidentalis is contrast-
ingly rounded, nonangulate laterally, and the in-
ner mandibular margin is nondentate (or at most
with a rudimentary tooth). The usual presence

of a foretarsal rake is an additional recognition
feature.

The female of occidentalis is less distinctive.
It is primarily characterized by the combination
of the impunctate mesopleuron, nonridged or
only finely ridged propodeal side, the suberect
setae on the hypoepimeral area, and the evenly
arcuate clypeal lip (neither emarginate mesally
nor indented laterally). Females of several other
species share these features: apricus, idiotri-
chus, some krombeini, mirandus, many semi-
rufus. Most of them have various prominent di-
agnostic features which are absent in
occidentalis. Furthermore, occidentalis and
some undescribed species have the clypeal free
margin less concave between the lobe and orbit
than do the other species. Subsidiary recogni-
tion feature of female occidentalis is the red
hindfemur (partly or all) and hindtibia.

GEOGRAPHIC DISTRIBUTION. — Xeric areas
west of the Rocky Mountains, north to Oregon
and Idaho, south to Arizona and Lower Califor-
nia.

MATERIAL EXAMINED. — HOLOTYPE: d , California, Inyo
Co., 12 mi. E Lone Pine, 19 May 1970, R. M. Bohart (UCD).

PARATYPES: 53 9 , 96 d , 4 Apr. to 23 July, 3 Aug., 28 Sep.
Specimens for which institution is not indicated below are all
in UCD.

UNITED STATES OF AMERICA

Arizona. Apache: Lukachukai, L. Burroughs, J. Bigelow,
MAC (1 9 , ASU). Coconino: 4.5 mi. E Moenkopi, J. H. & J.
M. Davidson, MAC (2 9 , ASU). Mohave: 8 mi. NE Mesquite
(in Nevada), FDP & PFT (19, USU). Navajo: Jadito Trade
Post, same collectors (2 9 , ASU).

California. Fresno: Jacolitos Canyon, RMB (1 d). Inyo:
Antelope Springs, MEI (19, UCR), 15 mi. s Big Pine, EEG
(I d); Deep Springs, D. Giuliani, MSW & J. S. Wasbauer,
MSW & J. Slansky (4 9, 13 d, CSDA; 1 d, WJP); same
locality, RMB, BV (6 d); Lone Pine, N. W. Frazier (1 9); 3
mi. N Lone Pine, JAP (I d , CIS); 7.3 mi. w Lone Pine, Ball-
mer & Bath(l 9); 12 mi. NE Lone Pine, RMB, EEG(1 9, 6
d). Kern: 14 mi. w Shatter, RMB (1 9). Lassen: Hallelujah
Junction, MEI (2 d , UCR). Los Angeles: 1 mi. w Little Rock,
E. G. & J. W. McSwain (Id). Mono: Chalfant, BV (1 d); 7
mi. sw Lee Vining, A. D. Telford (1 9). Monterey: Arroyo
Seco Camp, PT (1 d, USU). Plumas: Chilcoot, N. B. & W.
M. Elliott (Id, NYSU). Riverside: Anza, RMB (1 d). San
Bernardino: Cronise Valley, JAP (1 d); Cronise Wash 15 mi.
E Baker, WJP (Id, WJP); Four Corners, R. W. Thorp (1
9, CIS); Kramer Hills, PDH, G. A. Marsh, ROS (3 d; 1 d,
WJP); Yermo, collector unknown (1 9 , CAS). San Diego: Bor-
rego Valley, RMB (Id), WJP (1 9 , CAS; 1 d , WJP); Borrego
State Park, MSW & J. S. Wasbauer (19, CSDA); Scissors
Crossing, F. G. Andrews (19, UCR). San Luis Obispo: 10
mi. w Simmler, PDH (2 d , CIS).

Idaho. Cassia: 2.5 mi. s Malta, R. L. Westcott (19, UIM).
Franklin: Preston, GEB (Id. USU). Fremont: 6 mi. NW St.

NEW SPECIES OF NORTH AMERICAN TACHYSPHEX

Anthony, D. S. Horning (1 9,3d, UIM), R. L. Westcott (4
d, UIM); St. Anthony Sand Dunes, L. S. Hawkins (1 d,
UIM), R. L. Westcott (1 3, UIM), N. B. & W. M. Elliott (5
9 , 4 d , NYSU; 1 9 , 1 d , WJP). Lincoln: 4.5 mi. E Dietrich,
R. L. Westcott (1 9 , UIM); Shoshone, R. W. Haegele (1 S,
UIM).

Nevada. Churchill: 12 mi. NE Stillwater, FDP (3d; 1 9,2
S, USNM). Humboldt: Orovada, MEI (1 d); 10 mi. N Win-
nemucca, RMB (19). Lyon: Fernley, T. R. Haig (19); Yer-
ington, R. W. Lauderdale (1 9). Washoe: Nixon, RMB, R. J.
Gill, MEI, FDP, J. E. Slansky (89,4 d), MEI (1 9 , UCR),
R. L. Westcott (2 9 , 1 d , LACM); Patrick, FDP (1 9); 15 mi.
E Reno, RMB, MEI, FDP (9 d); Wadsworth, FDP (29,1
d); 2.8 mi. w Wadsworth, G. I. Stage (2 d , CAS).

Oregon. Harney: 21.5 mi. NW Fields sand dunes, R. L.
Westcott (1 9.OSDA).

Utah. Duchesne: 5.5 mi. w Roosevelt, R. W. Thorp (1 6",
CIS). Emery: Goblin Valley, FDP (2 9 , USU). Grand: 25 mi.
s Moab, GEB & R. Brumley (1 d; 3 d, USU; 1 d, WJP).
Millard: 6 mi. N Delta, S. M. Hogue (1 9, UIM); 15 mi. N
Delta, PFT (1<J, USU); 12 mi. NW Fillmore, FDP & Vincent
(2 d, USU). Washington: Santa Clara, GEB (19, USU).

Wyoming. Sweetwater: 20 mi. w Parson, PFT (1 d , USU).

MEXICO

Lower California. Punta de Cabras (12 mi. w of km 180 s
San Thomas), S. & S. Frommer (Id).

Tachysphex papago sp.n.

ETYMOLOGY. — Named after the Papago In-
dians of Arizona.

DIAGNOSIS. — Like psilocerus, papago has the
punctate mesopleuron, metapleuron, and pro-
podeal side (propodeal side impunctate in some
psilocerus), apicomesally unsculptured terga I
and II (smooth part contrasting with remaining
surface), and the largely brown or red female
flagellum. Unlike psilocerus, the upper meta-
pleuron of papago is simple, the metapleural
flange is narrow, the malar space is absent, and
the hindwing base is broad. The presence of
erect setae on the midfemoral venter in papago
(setae length about 1 DOA) is a subsidiary di-
agnostic feature.

GEOGRAPHIC DISTRIBUTION. — Southern Ari-
zona.

MATERIAL EXAMINED. — HOLOTYPE: 9, Arizona, Santa
Cruz Co., Nogales, Apr. 1937, R. C. L. Perkins (BMNH).

PARATYPES: Arizona, same data as holotype (Id, BMNH).
Cochise: 5 mi. w Portal, 7 May 1977, collector unknown (1
d, NYSU); 6 mi. w Portal, 7 & 12 May 1973, A. L. Steiner
(2 9 , UAE, WJP).

Tachysphex Solaris sp.n.

ETYMOLOGY. — The specific name solans is
Latin adjective meaning of the sun; it refers to
the sunny habitats in which the species occurs.

DIAGNOSIS. — Most solans can be recognized
by the uniformly yellowish humeral plate of the
forewing base (the median plate is usually con-
trastingly dark), but in some specimens the plate
is partly dark. In other species the humeral plate
is all dark or with a dark spot at the middle.
Subsidiary diagnostic features of Solaris are:
small size (body length 5-7.5 mm); clypeal free
margin shallowly concave between lobe and or-
bit; vestiture largely concealing mesopleural in-
tegument; setae oriented mainly transversely on
propodeal dorsum; and densely pubescent male
sterna.

GEOGRAPHIC DISTRIBUTION. — Xeric areas of
southern California and adjacent areas of Ne-
vada and Arizona.

MATERIAL EXAMINED. — HOLOTYPE: 9 , California, San
Diego Co., Borrego Valley, 20 Apr. 1957, R. C. Bechtel
(UCD).

PARATYPES: 59 9 , 26 d, 2 Apr. to 17 June.

Arizona. Mohave: 8 mi. E Mesquite (Nevada), FDP & PFT
(1 9, USU). Final: 5 mi. NW Coolidge, A. D. Telford (2 d,
UCD). Yuma: 6 mi. SE Parker, S. A. Gorodenski, JMD, MAC
(1 9 , ASU); 8 mi. SE Parker, J. H. & J. M. Davidson, MAC
(3 9 , UCD); 15 mi. E Yuma, PFT, FDP, GEB (19, USU);
18 mi. NE Yuma, FGW & GDB (1 d , UCD).

California. Imperial: Glamis, F. G. Andrews (19, CSDA;
1 9 , WJP); Pinto Wash, FXW (3 d , CAS); Palo Verde, RMB
(1 d,UCR), MSW(1 9, 1 d,CSDA; 1 d, WJP). Inyo: 13 mi.
s Death Valley Junction, LAS & ASM (Id, UCD); Lone
Pine, RMB (19, UCD); 2 mi. E Lone Pine, RMB (1 d, UCD),
15 mi. s Panamint Springs, PDH (2 9 , 2 d , UCD); s end
Owens Lake, M. S. & J. S. Wasbauer (3 9 , CSDA). Riverside:
18 mi. w Blythe, FDP (2 9, UCD), WJP (23 9, 3 d, CAS);
Hopkins Well, PDH (19, UCD); Thousand Palms, FDP (1
d, UCD). San Bernardino: Bagdad, JAP (2 d, CIS); Colton
Hills, TG (1 9 , TG), Cronise Valley, FDP (2 9 , UCD); Kelso,
N. J. Smith (1 9, UCD); 7 mi. sw Kelso, M. S. & J. S.
Wasbauer (1 d , CSDA); 10 mi. E Twenty nine Palms, collector
unknown (1 d, CSDA). San Diego: Borrego Valley, RCB,
JCH, H. R. Moffitt, EIS (6 9, 1 d, UCD; 2 9, USNM; 2
9 , 1 d , WJP); FXW (2 9 , 2 d , CAS).

Nevada. Clark: Glendale, FDP & PFT (1 9 , 1 d , USU); 20
mi. w Glendale, FDP & PFT (19, USU).

Tachysphex spatulifer sp.n.

ETYMOLOGY. — The specific name spatulifer
is a combination of the Latin word spatula, a
broad, flat tool for stirring or mixing, and the
suffix -fer, a bearer; with reference to the female
clypeus.

DIAGNOSIS. — The female of spatulifer has a
distinctive clypeus: the lip is broadened mesally,
usually variably, obtusely dentate (including one
median tooth). The clypeus is somewhat similar
in crenulatus, musciventris, arizonac, and la-

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3

mellatus, but unlike the former two species the
clypeal lip of spatulifer is not emarginate me-
sally, and unlike the latter two the middle scutal
setae are oriented posterad and the mesopleuron
is impunctate.

The male of spatulifer shares with crenulatus
the following diagnostic combination of char-
acters: clypeal lip triangular, middle scutal setae
oriented posterad, mesopleuron impunctate,
sterna evenly punctate, with nonvelvety pubes-
cence. Unlike crenulatus, the trochanteral
punctures of spatulifer are subcontiguous, the
lip corners are usually closer to the orbit than
to each other, and often the propodeal side is
coarsely ridged, the gastral apex is black, and
the frontal vestiture is golden.

GEOGRAPHIC DISTRIBUTION. — Washington to
California, east to southern Idaho and northern
Utah.

MATERIAL EXAMINED. — HOLOTYPE: 9, California, Mon-
terey Co., Arroyo Seco Camp, 11 May 1959, A. S. Menke
(UCD).

PARATYPES: 53 9, 85 8, 10 Apr. to 25 June (California,
Idaho, Oregon), 5-11 Aug. (Utah, Washington). Specimens
for which institution is not indicated below are kept in UCD.

California. Alameda: 1 mi. E Mission Peak, W. W. Middle-
kauf (I 9, 6 d; 1 3, CIS). Amador: Volcano, RMB (1 3).
Contra Costa: Mt. Diablo, RMB (4 6 ; 1 d , WJP), J. G. Rozen
(1 6), collector unknown (1 3; 1 3, USU); Las Trampas
Ridge (w Danville), FXW (1 9, 1 3, CAS). Eldorado: Pla-
cerville, GEB (1 9), D. J. Burdick (1 3 , CIS). Fresno: Deer
Cove Cr., TG (1 9, TG); Watts Valley, B. J. Adelson (1 9,
CIS). Kern: Glennville, RMB (2 9); Tejon Canyon, E. C.
VanDyke (1 9); 1 mi. E Woody, JAP, C. A. Toschi (2 3,
CIS). Lake: N. Fork Cache Cr. x Hwy. 20, MEI (13). Mar-
iposa: El Portal, RMB (Id); Indian Flat, RMB (19). Men-
docino: Mendocino, S. F. Cook (19, CIS). Monterey: Arroyo
Seco Camp, RCB, RMB, D. J. Burdick, HKC, C. Goodpas-
ture, FDP, LAS, PFT, BV (8 9 , 21 d ; 2 9 , 5 d , UCR; 2
9 , 5 d , USNM; 4 d , USU; 39,3d, WJP); Monterey, FDP
(19,3 d). Napa: Samuel Springs, RCB (1 d). Placer: 4 mi.
s Rocklin, MSW, P. Adams (1 9, 2 d, CSDA). Riverside:
Pinyon Flat, San Jacinto Mts., E. C. VanDyke (19). Sacra-
mento: Folsom, RMB (19), MSW (Id, CSDA); 10 mi. NE
Folsom, MSW (3 d); N Sacramento, PDH (7 d, CIS). San
Diego: Sorrento, JAP (1 9, CIS). San Luis Obispo: 2.5 mi.
Creston, C. A. Toschi (1 9 , CIS); La Panza Camp, 12 mi. NE
Pozo, JAP, R. W. Thorp (2 9 , CIS); Pozo, PDH, JAP, C. A.
Toschi (2 9 , 2 d , CIS); 3 mi. E Pozo, S. W. Thorp (Id, CIS);
5 mi. E Santa Margarita, W. S. & E. S. Ross (1 9, CAS).
Santa Clara: Mt. Hamilton, collector unknown (1 9); Los
Gatos, J. A. Kusche (19). Siskiyou: Windy Camp, TG (1
d, TG). Solano: Mix Canyon, R. B. & L. S. Kimsey (1 d).
Stanislaus: 3.2 mi. w Hwy. 120 on Evergreen Road, R. W.
Brooks (19). Tulare: Camp Wishon, TG (2 9 , TG); Sequoia
National Park, "Ash Mt. R.," EIS (1 d); Tule River Indian
Reservation, R. P. Allen (1 9, CAS); Wood Lake, N. W.
Frazier (19). Tuolumne: 4 mi. E Sonora, J. G. Rozen (19).

Yolo: Bear Cr. and Cache Cr. junction, ASM (19); Davis,
FDP, EIS (3 9, 1 d); Rumsey, ASM (19).

Idaho. Oneida: Black Pine Canyon, Malaise trap (19,1
d, USU).

Oregon. Umatilla: Athena, K. Gray & J. Schuh (1 9).

Utah. Cache: W. Hodges Canyon, Knowlton & Hanson (1
9, USU).

Washington. Pacific: Nahcotta, collector unknown (1 9,
CIS).

Tachysphex verticalis sp.n.

ETYMOLOGY. — The specific name verticalis is
derived from vertex, which is unusually narrow
in this species.

DIAGNOSIS. — Tachysphex verticalis resem-
bles many species of the brullii group (such as
belfragei or mundus) in having a longer than
wide vertex (as long as wide in some females)
and a fine thoracic sculpture. Unlike these
species, the propodeal dorsum setae of verti-
calis are inclined obliquely anterad, and the fe-
male tarsomere V is simple (apicoventral margin
not produced into a lobe, claws short, not pre-
hensile). T. verticalis differs from other species
of the pompiliformis group in having a uniformly
microareolate, impunctate mesopleuron and
propodeal side, and the contrastingly ridged pro-
podeal hindface. The markedly convex middle
clypeal section is also distinctive. Like crenu-
latus and glabrior, the lip of the female clypeus
has two lateral incisions on each side, but unlike
these species the vertex has a shiny, median sul-
cus which extends posterad from the postocellar
impression. The male differs from other species
of the group by its longer than wide vertex; like
glabrior, idiotrichus, and irregularis, sterna III-
VI have graduli.

GEOGRAPHIC DISTRIBUTION. — Northern
Mexico, southern Arizona and southwestern
New Mexico, California and adjacent areas of
northern Nevada; also isolated in Idaho and
Utah.

MATERIAL EXAMINED. — HOLOTYPE: 9, California, River-
side Co., 9 mi. w Beaumont, 29 July 1957, J. E. Gillaspy
(UCD).

PARATYPES: 78 9 , 165 d, 24 May to 5 Oct., mostly UCD,
but also AMNH, CAS, CIS, CSDA, HKT, KU, LACM,
MCZ, NYSU, UAT, UCR, UIM, USNM, WJP.

UNITED STATES OF AMERICA

Arizona. Cochise: Box Canyon (Chiricahua Mts.), RMB (1
9, 6 d); Portal, H. & M. Townes (1 d); Skelton Canyon, 6
mi. SE Apache, PDH (Id). Gila: Gila R. 3 mi. sw Christmas,
FGW (2 9, 1 d). Pima: Baboquivari Mts., O. C. Poling (1
d); Gates Pass, D. Graham (1 9); 2 mi. E Robles Pass, D.

NEW SPECIES OF NORTH AMERICAN TACHYSPHEX

Graham (2 9 , 1 d); Sabino Canyon, GEB, RMB, FDP, LAS
(29,5 d); Tanque Verde, FGW (1 d); Tucson, FDP, LAS
(2cJ).

California. .Mann-da: Tesla, J. E. Gillaspy (3 9). Butte: 1
mi. NE Pulga, R. L. Langston (2 d, CIS). Inyo: Antelope
Springs (8 mi. sw Deep Springs), RMB, HKC, T. H. Ganten-
bein, EEG, PMM, JAP (1 9 , 8 d , CAS; 1 6 , CIS; 19,10
<J), JAP, G. I. Stage (3 d, CIS); 3 mi. w Big Pine, LAS (1
9); Big Pine Cr., RMB, ROS (2 d); 10 mi. N Bishop, RMB
(1 d); Lone Pine Cr., RMB, EEG (2 9), Westgard Pass, H.
B. Leech (1 d, CAS); 4 mi. w Westgard Pass, H. V. Daly (1
9 , CIS). Kern: 2 mi. w Frazier Park, JAP (1 6, CIS); Kern-
ville, D. E. Hardy (1 d, KU). Lassen: Hallelujah Junction,
RMB (18). Los Angeles: Camp Baldy, W. A. McDonald (1
d, LACM); Crystal Lake Road, RMB (1 d); Elizabeth Can-
yon, EIS (1 6); La Crescenta, RMB (1 d); Mount Wilson
Road, RMB (19,1 d); Monrovia, R. Rosay (1 9); Sangus,
collector unknown (1 9, NYSU); Santa Susana Pass, MEI
(2 d); Tanbark Flat, RMB (3 d), FXW (1 9, 2 d, CAS).
Mariposa: Jerseydale, P. D. Levin (1 d, UIM). Monterey: Mill
Cr. (Santa Lucia Mts.), EIS (1 d); Paraiso Springs, L. S.
Slevin (2 9); 4 mi. s San Ardo, P. E. Paige (2 6); 4 mi. w
Soledad, G. R. Ballmer (19, UCR). Orange: Upper Trabuco
Canyon, E. M. Fisher (19). Placer: Lake Tahoe, collector
unknown (19). Riverside: Anza, RCB (19); Banning, JCH
(1 9); 9 mi. w Beaumont, J. E. Gillaspy, JCH, H. R. Moffitt
(3 9 , 9 cJ); Pinyon Flat, P. H. Arnaud (1 6 , CAS); Riverside,
JCH (2 9 , UCR); San Timoteo Canyon, MSW, R. McMaster,
A. Hardy, J. Slansky (5 9 , 12 d); Whitewater, M. J. Stebbins
(1 d). San Bernardino: Cajon, RCB, RMB, H. R. Moffitt (5
d); Cajon Junction, D. Burnett (1 9); Hole-in-the-Wall (Prov-
idence Mts.), TG (1 d , TG); Mill Cr. Canyon, JCH (2 9); Oak
Glen, TG (1 9 , 1 d , TG); Upper Santa Ana R., A. L. Melan-
der (19); Wildwood Canyon, H. R. Moffitt (1 d); 3 mi. SE
Yucaipa, TG (1 d , TG). San Diego: Laguna Mts. road, FXW
(1 d , CAS); La Jolla, K. Corwin, P. Adams (1 9 , 2 d), J. C.
Bridwell (1 d); Julian, H. R. Moffitt (2 d), FXW (1 d, CAS);
Poway, F. Blaisdell (1 d); San Diego, H. E. & M. A. Evans
(2 9), H. A. Hill (Id); Scissors Crossing, EIS (Id); Sorren-
to, JAP (19, CIS); 9 mi. s Warner Springs, RMB (3 d). San
LuisObispo: Creston, L. E. Guenther(l d); Nacimiento Dam,
JAP (1 9, CIS); 3 mi. NW Paso Robles, R. L. Langston (1
9 , CIS). Santa Barbara: Bluff Camp (San Rafael Mts.), PMM,
FDP (3 d); 3 mi. w Cachuma Lake, RMB, P. E. Paige, FDP,
J. R. Russel (4 d), W. A. Steffan (2 9 , CIS); Los Prietos, J.
S. Buckett (Id), JAP (Id, CIS); Santa Ynez Mts., RMB,
PMM, ASM, FDP (6 d). Santa Clara: San Antonio Valley,
3.5 mi. N Del Puerto Canyon road, JAP (1 9 , CIS). Shasta:
Hat Cr. P.O., E. E. Lindquist (1 9 , CIS); Redding, T. R. Haig
(1 d). Stanislaus: Del Puerto Canyon, RMB, N. J. Smith (3
9, 3 d), BV (3 9, 4 d). Trinity: Junction City, T. R. Haig
(1 9, 1 d), 3 mi. w Weaverville, DRM (1 d, UCD). Tulare:
Three Rivers, H. R. Moffitt (Id, UCR). Ventura: Foster
Park, RMB, ASM (2 d); Sespe Canyon, PMM, FDP, RMB
(3 d), W. A. Steffan (1 d, CIS). Also: County unknown, Oak
Grove, J. Wilcox (Id, OSU).

Idaho. Owyhee: 2 mi. sw Murphy, D. S. Horning, DRM (1
(J).

Nevada. Douglas: 3 mi. s Genoa, RMB (1 9,3d); Minden,
RCB, RMB (7 9 , 1 d). Storey: Geiger Summit, W. H. Lange
(Id). Washoe: 54 mi. NW Gerlach, FDP (4 d).

New Mexico. Hidalgo: Granite Gap, 18 mi. N Rodeo, RMB
(29,3d, AMNH; 3 9 , 2 d , UCD).

Utah. Grand: Moab, GEB (19). Washington: Leeds Can-
yon, G. F. Knowlton (19, USU).

MEXICO

Lower California. 10 mi. E Bahia San Quintin, FXW (1 9 ,
CAS); 4 mi. s La Rumorosa, MEI (1 9, 1 d, UCR); 4 mi.
wsw Miraflores, J. Slansky, M. K. & C. Wasbauer (19,2
d); San Quintin, FXW (Id, CAS).

Sonora. Cocorit, FDP(1 d, UCD).

Tachysphex yolo sp.n.

ETYMOLOGY. — Named after Yolo County,
California, where the holotype was collected.

DIAGNOSIS. — Tachysphex yolo is character-
ized by the transversely oriented middle scutal
setae, sternum I with a horizontal depression at
apex, and the usually punctate mesopleuron
(punctures indistinct in some individuals). Other
species share this combination of characters, but
the female of yolo has a distinctive clypeus and
sternum II: the dense clypeal punctation attains
the lip base laterally, so that the sparsely punc-
tate, apical area does not extend laterad to the
lip corner level (its width is about 0.5-0.8 of the
lip foremargin); the micropunctation of sternum
II is absent along the midline from the base (or
near base) to apex. In the other species, the
dense punctation does not attain the lip base,
and the sparsely punctate, apical area is as wide
as the lip or nearly so; and the micropunctation
of sternum II is usually absent only from an ap-
icomedian, triangular area. The male resembles
lamellatus and sonorensis in having nonvelvety
sternal pubescence, but unlike these species its
clypeal lip is not triangular. Unlike most lamel-
latus, the hypostomal carina of yolo is not la-
melliform, and unlike the male of sonorensis,
the flagellomeres III and IV are about equal in
length. Subsidiary recognition features are: me-
sopleural punctures subcontiguous, ridges of
propodeal side evanescent in many specimens.

MATERIAL EXAMINED. — HOLOTYPE: d, California, Yolo
Co., Davis, 4 June 1961, F. D. Parker (UCD).

PARATYPES: 312 9 , 414 d , 1 Apr. to 14 Oct. Specimens for
which institution is not indicated are all in UCD.

UNITED STATES OF AMERICA

Arizona. Cochise: Bowie, A. D. Telford (3 9); Willcox,
RMB, PDH, D. D. Linsdale (2 9, CIS; 10 9, 4 d; 2 9,
USNM); 1 mi. s Willcox, FGW, E. Erickson (1 9 , UAT); 3.5
mi. s Willcox, E. G. Linsley (19, CIS). Coconino: Moenkopi,
RMB(1 9); 3 mi. SE Moenkopi, ASM & WJP(1 d, WJP); 4.5
mi. E Moenkopi, J. M. & M. A. Davidson & MAC (3 9,
ASU). Graham: s side of San Carlos Reservoir, D. & J. Schuh
(Id, OSU). Maricopa: Gila Bend, GDB (2 9); 5 mi. w Gila

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3

Bend, GDB & FGW (1 9); 20 mi. s Gila Bend, R. H. & E.
M. Painter (Id, UAT); Sentinel, GDB (3 9). Mohave: 4 mi.
w Chloride, GEB, PFT, FDP (3 9 , 5 d , USU); 9 mi. E Oat-
man, same collectors (5 9, 1 d, USU); 16 mi. N Wikieup,
FGW & GDB (Id). Navajo: Jadito Trade Post, J. H. & J. M.
Davidson & MAC (19, ASU). Pirna: Tucson, GDB (29,1
d, UAT), G. & A. Ferguson (1 9), R. X. Schick (1 9,
LACM). Pinal: 3 mi. w Oracle, FDW, Bequaert, Holler (1
d , UAT); Sacaton, A. D. Telford (19,1 d). Yavapai: 8 mi.
N Aguila, FDP (19, USU). Yuma: E Aztec, FGW & GDB
(2 9); Dateland, GDB & FGW (1 d); Parker, J. M. & J. H.
Davison (1 9, 2 d); 8 & 18 mi. SE Parker, J. H. & J. M.
Davidson & MAC (3 9 , ASU); 8 mi. SE Parker, S. A. Goro-
denski & JMD (Id, ASU), J. Heddock (19, CIS); 12 mi. s
Parker, MAC & M. A. Mortenson (1 9 , CIS); NW Vicksburg,
FGW & GDB (29,1 d); Yuma, FGW & GDB (1 d); 15 mi.
E Yuma, PFT, FDP, GEB (19, USU); 21 mi. N Yuma, MEI

d 9).

California. Alameda: Arroyo Valle, W. J. Turner (1 9 , CIS).
Colusa: 2 mi. E Colusa, MSW (1 9). Eldorado: Chile Bar, L.
W. Quate (1 9). Fresno: 25 mi. E Fresno, PFT (2 d, USU).
Imperial: Chocolate Mts., Ogilby road, 3 mi. s junction Hwy.
78, MSW (9 9 , CSDA); Glamis, FDP (Id); 20 mi. E Glamis,
FDP (1 9); Palo Verde, RMB, EEC, ROS (39,2 d), MSW
(1 9, 1 d, CSDA); 3 mi. s Palo Verde, C. A. Toschi (1 9,
CIS); Pinto Flat, FXW (19, CAS); Pinto Wash, FXW (Id,
CAS); 20 mi. w Yuma, PFT & FDP (19, USU). Inyo: An-
telope Springs (8 mi. sw Deep Springs), RMB, HKC, T. H.
Gantenbein (69,3d;69,lld, CAS), PDH (69, 13 d ,
CIS), JWMS (19, CIS), JAP (Id, CIS); Big Pine, C. D.
Michener (1 d, KU); Darwin Falls, RMB, EEG (19,1 d);
3.5 mi. s Death Valley Junction, N. L. Rump (1 9); 13 mi. s
Death Valley Junction, LAS (1 d); Deep Springs, RMB, BV
(29,3d), MSW, MSW & J. Slansky (2 9, 13 d, CSDA; 1
d, WJP); Eureka Valley, D. Giuliani (1 9, CSDA); 1 mi. N
Lone Pine, MEI (1 9 , 1 d , UCR); 2 mi. E Lone Pine, RMB,
EEG (29,9 d); Owens Lake, MSW & F. G. Andrews (2
9 , CSDA); Panamint Springs, PMM (19), FDP (Id); West-
gard Pass, H. V. Daly (Id, CIS); Wyman Canyon (White
Mts.), HKC (Id, CAS). Kern: Johannesburg, EIS (19);
Kernville, T. R. Haig (1 9). Lake: N. Fork Cache Cr. x Hwy.
20, D. Q. Cavagnaro (19). Lassen: Hallelujah Junction, MEI
(2 d, UCR). Los Angeles: Huntington Park, A. Bauman (1
9); 8 mi. N Llano, JCH, EIS (3 9). Mendocino: Navarro, N.
B. & W. M. Elliott (4 d, NYSU); Navarro R. x Hwy. 128,
Hendy Groves State Park, MSW (99, 13d, CSDA; 19,2
d , WJP); Robinson Cr. (4 air mi. sw Ukiah), TG (3 d , TG).
Monterey: Fort Ord, HKC (Id, UCD); Monterey, FDP, L.
S. Slevin (3 d); Soledad, RMB (3 d). Modoc: Adin Pass, T.
R. Haig (1 9, CSDA). Mono: Benton Inspection Sta., RMB
(1 d); Paradise Camp, FDP (1 d). Plumas: Chilcoot, N. B.
& W. M. Elliott (1 9 , 5 d , NYSU); Halsted Campground (E.
Branch N. Fork Feather R.), P. H. Arnaud (1 9 , 1 d, CAS).
Riverside: Andreas Canyon, RMB (2 d); Anza, RMB (29,2
d); 8 mi. E Banning, R. R. Snelling (Id, LACM); 9 mi. w
Blythe, JWMS (19, CIS); 18 mi. w Blythe, RMB, JCH, D.
S. Horning, FDP (1 9 , 4 d ; 2 9 , WJP), WJP (Id, CAS); 5
mi. N Desert Center, C. D. MacNeill (2 9 , CIS); 17 mi. E
Desert Center, Rosen & Schrammel (1 9); 5 mi. s Hemet,
RMB(1 9);Indio, PDH(1 d), MEI(1 9 , UCR); Joshua Tree
National Monument, PFT (2 d , USU); Palm Springs, JWMS
(1 9); 2 mi. E Palm Springs, EIS (19); Riverside, JCH (1
d), EIS ( 1 d , UCR); Temecula, EIS (Id); Thousand Palms,
RMB, EEG, H. R. Moffitt, FDP (39,9 d), W. R. Richards

(1 9, CNC); Whitewater, JWMS (1 9); Wiley Well, RMB (I
d). Sacramento: Grand I., MSW (1 9, CSDA); Sacramento,
RMB, FDP (3 d), MSW (4 d, CSDA); Sacramento (Sacra-
mento R. Levee), MSW (Id), MSW & F. G. Andrews (2
d, CSDA). San Benito: Pinnacles, PDH (Id, CIS). San Ber-
nardino: 1 mi. s Adelanto, MEI (1 d); 10 mi. s Adelanto, J.
A. Froebe(l d); Bagdad, JAP (1 d,CIS);4& 14 mi. s Baker,
M. S. & J. S. Wasbauer (2 d , CSDA); Cottonwood Wash, TG
(2 9 , TG); Cronise Valley, FDP (1 9); Cronise Wash (15 mi.
E Baker), WJP (2 9 , 5 d , CAS); 12 mi. SE Ivanpah, PDH (1
9, 1 d); Joshua Tree National Monument, TG (1 d, TG);
Kelso Dunes, TG (4 9 , TG); Kelso Mts., TG (1 d, TG); Kra-
mer Hills, G. A. Marsh (1 d , CIS), ROS (1 9); 3 mi. s Kramer
Junction, MEI (2 d); 14 mi. s Kramer Junction. JWMS (1
9); 20 mi. w Landers, PFT & N. Youssef (1 d , USU); 12 mi.
ESE Tecopa, MSW, T. Eichlin (1 d, CSDA); 22 mi. N Manix,
G. A. Marsh (1 9); 2 mi. w Phelan, EIS (2 d); 14 mi. w Rice,
C. D. MacNeill (1 d, CIS); Twentynine Palms, R. R. Pinger
(Id, CSDA); 32 road mi. E Twentynine Palms, TG (1 9, 1
d, TG); Vanwinkle Spring, G. E. Wallace (I d); 11 mi. E
Yermo, JWMS (19, CIS). San Diego: 8 mi. E Banner, JAP
(2 d, CIS); Borrego Valley, RMB, EEG, PDH, FDP, EIS,
MSW ( 19 9 , 29 d ; 2 d , WJP), MEI (2 d , UCR), G. A. Marsh
(1 9, CIS), MSW, J. Slansky, Adams (1 9, 4 d, CSDA),
MSW (1 9 , 4 d , CIS), FXW (2 9 , 29 d , CAS); Del Mar, C.
H. Frady (19, OSU); 1 mi. s Del Mar, PDH (2 9, CIS);
between Ocotillo & Borrego, A. R. Moldenke (2 d , LACM);
Scissors Crossing, EIS (2 9), MSW (19, CSDA); Sorrento,
JAP (1 9 , 7 d , CIS); 2 mi. N Warner Springs, B. M. Bartosh,
RCB (3 9 , 2 d). San Luis Obispo: Black Lake Canyon, RMB
(19,1 d); 10 mi. w Simmler, PDH (2 d, CIS). San Mateo:
10 mi. sw San Francisco, W. Bohart (Id, USU). Santa Bar-
bara: 3 mi. w Cachuma Lake, P. E. Paige (19), Los Prietos,
J. S. Buckett (1 d), JAP (1 9, CIS); 2 mi. E Solvang, JAP
(19, CIS). Santa Clara: San Jose, PFT (19, USU). Siskiyou:
between Hawkinsville & Lona Gulch, BV (1 9 , CSDA). Sut-
ler: Nicolaus, MSW (1 9, 17 d, CSDA). Trinity: Hayfork
Agricultural Inspection Sta., JAP (2 d, CIS); Junction City,
T. R. Haig (2 9 , 1 d , CSDA). Ventura: Foster Park, J. L.
Bath (19, UCR), J. R. Russell (19); Ventura, J. R. Russell
(1 9). Yolo: Capay, R. E. Rice (1 9); Davis, RMB, C. G.
Moore, C. R. Kovacic, ASM, L. R. Nault, FDP, WJP, LAS
(12 9, 28 d; 2 d, USNM; 29,3d, WJP); 3 mi. s Davis, R.
R. Snelling (1 d, LACM); Putah Canyon, FDP (19,2 d);
Rumsay, RMB (1 d), AMS & LAS (1 9, 1 d, LACM).

Idaho, Franklin: Preston (19, USU). Fremont: St. Anthony
Sand Dunes, N. B. & W. M. Elliott (1 9 , 1 d, NYSU). Lin-
coln: 6 mi. NE Shoshone, A. R. Gittins (d , UIM).

Nevada. Churchill: 23 mi. E Fallen, E. G. Linsley (1 9); 3
mi. w Hazen, ASM (1 9), 4 mi. E Hazen, MEI (Id, UCR);
Sand Mt. (9 mi. Frenchman), J. Doyen (1 9, CIS). Clark:
Jean, GEB (1 d); 9 mi. sw Mesquite, RCB (10 9, 6 d); 30
mi. s Searchlight, PFT, Rust, Youssef (1 d, USU); Valley of
Fire, PFT, FDP, GEB (Id, USU). Humboldt: 10 mi. N Win-
nemucca, EEG (1 9). Lyon: Weeks, FDP (1 9). Mineral: Lu-
ning, R. F. Denno & DRM (2 d); 3 mi. SE Schurz, FDP (1
d). Pershing: Woolsey, T. R. Haig (2 9, CSDA). Washoe:
Nixon, RMB, FDP (1 9,4d),MEI(l d, UCR); Patrick, FDP
(1 <?)•

New Mexico. Dona Ana: Las Cruces, RMB (1 9, 2 d), R.
H. Beamer(l 9, KU); 4 mi. E Mesilla Park, PDH (1 d, CIS).
Lincoln: 5 mi. s Oscuro, R. L. Westcott (19, UIM). Otero:
Alamogordo, collector unknown (19, CU); White Sands Na-
tional Monument, H. V. Weems (1 9, FSCA). Socorro: La

NEW SPECIES OF NORTH AMERICAN TACHYSPHEX

Joya, 20 mi. N Socorro, H. E. Evans (6 2,3 <J, CSU).
Valencia: Acoma Pueblo, ASM & WJP (1 $ , WJP).

Oregon. Destitutes: Smith Rock State Park, M. B. & W. M.
Elliott (8 9, 15 6, NYSU). Josephine: 8 mi. w Grants Pass,
R. L. Westcott (1 9, ODA). Klamath: Bonanza, J. Schuh (1
9). Morrow: Boardman, G. R. Ferguson (2 d). Umatilla: Hat
Rock State Park, EIS (19).

Texas. Hudspeth: McNary, H. E. Evans (2 9 , 1 d , CSU);
Sierra Blanca, RMB (19). Presidio: 5 mi. E Presidio, D. S.
Horning (Id).

Utah. Cache: Cornish, GEB & PFT (5 9 , 2 6 , USU; 2 9 ,
WJP). Emery: Goblin Valley, FDP (1 d, USU), 2 air mi. w
Little Gilson Butte, TG (1 d, USU). Juab: 12 mi. s Eureka,
JWMS (19, CIS); White Sand Dunes (25 mi. sw Eureka), W.
F. Barr(l 3, UIM). Millard: 15 mi. N Delta, PFT (1 d,USU);
Pahvant (near Flovell), GEB & E. A. Cross (Id, UCD). San
Juan: 6 mi. s La Sal Junction, R. W. Thorp (1 d); 25 mi. s
Moab, GEB, R. Brumley (2 9 , 1 d; 1 9 , USU). Washington:
Leeds Canyon, G. F. Knowlton, W. J. Hanson, T. H. Hsiao
(1 9, 2 d, USU).

MEXICO

Lower California. 10 mi. E Bahia San Quintin, FXW (1 d,
CAS); Descanso, RMB (1 9); El Pescadero, MSW, J. Slansky
(3 9 , 6 d, CSDA); La Paz, FXW (2 d, CAS); Los Barriles,
MSW (3 9, CSDA); 20 mi. N Mesquital, Ross & Bohart (1
2); Progreso, Sierra Juarez, FXW (1 9,3d, CAS); 38 km s
Rosarito (114°), E. Fischer, R. Westcott (1 9 , CAS); San Car-
los, H. E. Evans, W. Rubink & D. Gwynne (4 9 , CSU); 3
mi. N San Felipe, MEI (3 9 , 1 d , UCR); 15 mi. N San Ignacio,
Ross & Bohart (1 9); 10 mi. s San Quintin, J. Slansky, M.
& K. Wasbauer (1 9, CSDA); San Vicente, C. H. Frady (1
9,OSU), JAP(1 9, CIS).

Chihuahua. Moctezuma, JWMS (Id, CIS); Samalayuca,
RMB (2 9).

Sinaloa. 8 mi. s Elota, LAS (Id).

Sonora. Cerro Pinacate, McDougal Crater, GDB (1 d,
UAT); Guaymas, E. P. VanDuzee (1 9); 39 mi. s Puerto Pe-
nasco, M. Leppla, J. Bigelow, MAC, J. Davidson (1 9 , ASU);
"Sonora, 85 km. so.," A. L. Melander (19,6 d); Tepoca
Bay, E. P. VanDuzee (1 9).

Tachysphex yuma sp.n.

ETYMOLOGY. — Named after the Yuma Indi-
ans of Arizona.

DIAGNOSIS. — Tachysphex yuma is character-
ized by the punctate mesopleuron (punctures
shallow), middle scutal setae oriented postero-
laterad or (some males) posterad, and sternum
I with an apical depression. Several other
species share this combination of characters, but
they have a uniformly ridged propodeal side
(ridges evanescent in many yolo). In yuma, the
propodeal side is either coarsely ridged poste-
riorly and microridged along the metapleural
sulcus; or (most specimens) nonridged along the
metapleural sulcus and ridged along the dorsal
margin (or dorsal and posterior); or (some
males) all nonridged. Furthermore, the flagellum

of yuma is somewhat longer; for example, the
length of flagellomere IV is 3.6-4.2 (female) and
2.0-2.4 (male) times its width, and up to 3.2 and
2.0 times, respectively, in the other species. In
the male, the unusually broad clypeal lobe is
distinctive (corners markedly closer to orbits
than to each other) and the velvety sternal pu-
bescence is an additional recognition feature.

GEOGRAPHIC DISTRIBUTION. — Idaho, Ore-
gon, California, Arizona, southern Texas, north-
ern Mexico (Sonora, Lower California).

MATERIAL EXAMINED. — HOLOTYPE: d, Mexico, Baja Cal-
ifornia Sur, La Paz, 10-12 Oct. 1954, F. X. Williams (CAS
Type No. 13966).

PARATYPES: 8 9, 25 d, 2 Mar. (Sonora), 8 Apr. to 1 June,
14 July (Oregon), 9-15 Oct., 1 1 Nov.

UNITED STATES OF AMERICA

Arizona. Cochise: Canelo, A. & H. Dietrich (1 9, NYSU);
5 mi. w Portal, collector unknown (2 d, NYSU); 6 mi. w
Portal, A. L. Steiner (2 d, UAE, WJP). Maricopa: Wicken-
burg, PFT & GEB (Id, WJP). Coconino: Grand Canyon Na-
tional Park, 15 mi. NE Phantom Ranch, J. E. Slansky (1 d,
UCD).

California. Imperial: 20 mi. E Glamis, FDP (Id, UCD).
Inyo: Darwin Falls, ASM (1 d , UCD); Panamint Springs, MEI
(Id, UCD); Surprise Canyon, FDP (2 d, UCD, WJP). Riv-
erside: Boyd Desert Research Center, 4 mi. s Palm Desert,
PDH (1 9, 1 d, CIS; 1 9, WJP); Deep Canyon, MEI (3 d,
UCD, UCR, WJP), EIS (1 d, UCD); Salton Beach, A. L.
Melander (1 2, UCD); Thousand Palms Canyon, RMB (1
d, UCD); Whitewater, D. J. R. (1 d, CIS). San Bernardino:
3 mi. N Crossroads, C. D. MacNeill (1 d, CAS). Shasta: Red-
ding, DRM (19, UCD). Stanislaus: Empire, E. I. Beamer (1
9, KU).

Idaho. Twin Falls: Rock Cr. Canyon (19 mi. s Hansen), R.
L. Westcott (1 9, UIM).

Oregon. Malheur: 4 mi. N Juntura, H. A. Scullen (1 d,
UCD).

Texas. Brewster: 20 mi. NNW Marathon, M. Masters (1 d,
CU).

MEXICO

Baja California Norte. 65 mi. s San Felipe, R. D. Gehring
(1 d,CIS).

Baja California Sur. 3 mi. s Ignacio, MEI (2 d , UCR, WJP),
Rancho El Cayote, Maynard & Honey (Id, LACM).

Sonora. Bahia San Carlos, P. H. Arnaud (19, CAS).

SPECIES OF THE BRULLII GROUP
Tachysphex acanthophorus sp.n.

ETYMOLOGY. — The specific name acantho-
phorus is derived from the Greek words acan-
thos, a spine, andphorein, to bear, a spine-bear-
er; with reference to the tarsal spines.

DIAGNOSIS. — The female of acanthophorus
has a distinctive tarsal feature: one or two sub-
apical spines on each lateral margin of tarsomere

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3

V, and usually central spines on its venter; un-
like armatus, the tarsomere V lacks basoventral
spines. The lateral, subapical spines on tarso-
mere V occur also in most western females of
mundus, but unlike that species the mesopleural
vestiture is dense (partly obscuring sculpture) in
acanthophorus. Tarsomere V is similar in acan-
thophorus and the South American species spi-
nulosus, but in the latter the gaster is black and
the mid- and hindfemora are red.

The males of acanthophorus and armatus dif-
fer from other species of the brullii group in
lacking a clypeal bevel and graduli, and in hav-
ing a very narrow clypeal lip and a transverse
sulcus on sterna III-VI (the sulcus is visible only
when segments are fully extended). Unlike ar-
matus (in which the sternal surface is flat), ster-
num II of acanthophorus is somewhat swollen
along the foremargin of the apical depression.
Furthermore, the mesopleural sculpture is ob-
scured by vestiture or hardly visible in acantho-
phorus, but easily visible in armatus.

GEOGRAPHIC DISTRIBUTION. — Xeric areas of
southwestern U.S. and northern Mexico.

MATERIAL EXAMINED. — HOLOTYPE: 9 , Arizona, Cochise
Co., Willcox, 14 Aug. 1958, P. D. Hurd (UCD).

PARATYPES: 134 2 , 180 6 , 1 intersex; May to 7 Nov. Spec-
imens for which institution is not indicated are all in UCD.

UNITED STATES OF AMERICA

Arizona. No specific locality (1 9). Cochise: 1 mi. E Apache,
J. G. Ehrenberg (2 d, AMNH); Bowie, A. D. Telford (1 9,
WJP); 7 mi. SE Dos Cabezas, D. S. Chandler (1 d, UAT);
Douglas, G. L. Ballmer (2 9); 2 mi. E Douglas, R. W. Brooks
(1 9); Portal, K. V. Krombein (28 9, 22 3; BMNH, KVK,
WJP); 2 mi. E Portal, J. H. Puckle, M. A. Mortenson & MAC
(2 d); 2 mi. NE Portal, M. & T. M. Favreau (1 d, AMNH);
Willcox, RMB (2 9 , 2 d), D. D. Linsdale (1 9 , 5 d ; AMNH,
UCD), PDH (1 9, 2 d, CIS; 1 9, USNM). Maricopa: 5 mi.
N Aguila, GEB & PFT (19, USU); Tempe, collector un-
known (1 d, MCZ); 5 mi. SE Wickenburg, PDH (1 9, CIS).
Pima: 30 mi. SE Ajo, C. R. Kovacic (19); Continental, MEI
(2 d); Sabino Canyon (Santa Catalina Mts.), GDB & FGW
(Id, UAT), FDP & LAS (19); Tucson, C. L. Crow (19),
M. L. Lindsay (Id, UAT), FDP (Id), FDP & LAS (19).
Final: 5 mi. NW Coolidge, A. D. Telford (1 d); Picacho Pass,
DRM & J. E. Lauck (19); collector unknown (19, UCR);
Superior (Boyce Thompson Arboretum), GDB (Id, UAT).

California. Imperial: Brawley, GEB (1 9). Inyo: Antelope
Springs (8 mi. sw Deep Springs), HK.C (13 9 , 11 d ; AMNH,
UCD), PMM (3 9), DRM (19,1 d), JAP, G. I. Stage (11
9, 48 d, CIS); Deep Springs, H. Nakakihara (19, UCR).
Riverside: 12 mi. N Blythe (also 18 mi. w), RMB (2 9); 20 mi.
w Blythe, JWMS (3 9, 1 d, CIS); Indio, MEI (Id, UCR);
3.5 mi. s Palm Desert, S. Frommer & B. Morley (1 9, 1 d,
UCR). San Bernardino: no specific locality, D. W. Coquillett
(Id, USNM). San Diego: Borrego, A. L. Melander (19).

Tulare: Lemon Cove, J. C. Bradley (3 9 , NYSU; 4 d , CU);
Three Rivers, collector unknown (Id, CU).

Colorado: Bent: Hasty, H. E. Evans (9 9 , 2 d ; BMNH,
CSU, WJP).

Nevada. Clark: Sandy, RCB (Id). Mineral: Luning, R. F.
Denno & DRM (19). Nye: Mercury (Id, USNM).

New Mexico. Dona Ana: Las Cruces, RMB (19). Eddy:
15.5 mi. w Artesia, V. E. Romney (1 d, USNM). Grant: 25
mi. E Lordsburg, H. A. Scullen (1 d). Hidalgo: 21 mi. s An-
imas, J. G. & B. L. Rozen (19, AMNH); 22 mi. s Animas,
J. Rozen & M. Favreau (Id, AMNH); Cienaga Lake, J. H.
& J. M. Davidson & MAC (2 d); Granite Gap (17 mi. N Ro-
deo), F. G. Andrews (19, CSDA); Granite Pass area (20 mi.
N Rodeo), Hwy. 80, MSW, J. Slansky & C. Freeberg (19,
CSDA); 1 mi. N Rodeo, J. H. Puckle, M. A. Mortenson &
MAC (1 d); 4 mi. sw Rodeo, J. G. Rozen (19, AMNH).
Otero: White Sands National Monument, RMB (Id). Socorro:
La Joya (20 mi. N Socorro), W. Rubink (19, CSU; 2 d,
USNM). Quay: Tucumcari, RMB (2 d).

Texas. Bexar: no specific locality, H. B. Parks (3 9 ; UCD,
WJP). Brewster: Big Bend National Park (Nine Point Draw),
W. R. M. Mason (4 9 , 22 d ; BMNH, CNC, WJP); Big Bend
National Park (Boquillas), W. R. M. Mason (1 intersex, CNC);
Glenn Spring, F. M. Gaige (1 d). El Paso: Sierra Blanca, col-
lector unknown (1 d, CU). Hudspeth: Fort Hancock, CU
Exped. (19, NYSU); McNary, H. E. Evans (1 9, 1 d;
BMNH, CSU). Presidio: 3 mi. E Presidio, H. E. Evans (2
9, MCZ), J. E. Gillaspy (1 9).

Utah. Garfleld: Shootaring Canyon, D. Vogt (1 9, 1 d,
USU). Washington: Leeds Canyon, G. F. Knowlton, W.
J. Hanson, T. H. Hsiao (2 9 , 1 d , USU); St. George, GEB
(1 d); Toquerville, G. F. Knowlton, W. J. Hanson, T. H.
Hsiao (2 d , USU).

MEXICO

Baja California Sur. 4 mi. wsw Miraflores, J. Slansky, M.
K. & C. Wasbauer (1 9, 4 d, CSDA).

Chihuahua. 15 mi. s Chihuahua, H. E. Evans (2 9, 4 d;
BMNH, MCZ, WJP).

Sinaloa. Culiacan, H. E. Evans (Id, CU); s Lorenzo, GEB
& RMB (2 9 , USU); Mazatlan, W. R. M. Mason (1 9 , CNC);
2.5 mi. N Mazatlan, MSW (1 9 , CIS); 5 mi. N Mazatlan, MSW
& J. Chemsak (Id, CIS). Sonora. Alamos, RMB (1 d), W.
J. Hanson & T. L. Whitworth(l d.USU); 10 mi. sw Alamos,
FDP & LAS (Id); Desemboque, C. & P. Vaurie (Id); 19.4
& 20 mi. s Estacion Llano, MEI, EIS, P. A. Rauch (3 d,
UCR); Guaymas, RMB (1 9); 5 mi. s Magdalena, FDP & LAS
(1 d); Minas Nuevas, C. & P. Vaurie (1 d); 10 mi. E Navajoa,
W. L. Nutting & FGW (2 d , UAT); San Carlos, RMB (19,
1 d).

Tachysphex armatus sp.n.

ETYMOLOGY. — The specific name armatus is
a Latin word for armed, with reference to the
ventral spines of female hindtarsomere V.

DIAGNOSIS. — The female of armatus is unique
among the North American Tachysphex in hav-
ing basoventral spines on the tarsomere V. Oth-
erwise it is very similar to acanthophorus with
which it shares other structures of the tarsomere
V: one to several medioventral spines and one

NEW SPECIES OF NORTH AMERICAN TACHYSPHEX

or two preapical spines on each lateral margin.
The lateral spines are also found in most western
specimens of mundus.

The male of armatus is very similar to acan-
thophorus. See that species for differences.

GEOGRAPHIC DISTRIBUTION. — Desert areas
between southwestern Texas and southern Cal-
ifornia, also Lower California.

MATERIAL EXAMINED. — HOLOTYPE: 9 , Nevada, Clark
Co., Sandy, 24 July 1958, R. C. Bechtel (UCD).
PARATYPES: 17 9 , 9 3 ; May to 5 Sep.

UNITED STATES OF AMERICA

Arizona. Maricopa: 10 mi. E Gila Bend, GDB (19, UCD);
Phoenix, R. H. Crandall (1 ?, UCD); 5 mi. SE Wickenburg,
PDH (1 $ , CIS), P. H. Timberlake (1 6 , UCR). Pima: Tucson,
F. M. Carpenter (19, UCD).

California. San Diego: San Diego, F. E. Blaisdell (19,
UCD).

Nevada. Clark: Sandy, RGB (1 <S , WJP). Lincoln: Alamo,
FDP(1 9, WJP).

Texas. Brewster: Big Bend National Park, R. W. Strandt-
mann (1 9 , 1 6 ; USNM, UCD); Big Bend National Park (Nine
Point Draw), W. R. M. Mason (1 9, CNC). Hudspeth:
McNary, H. E. Evans (29,3d; CSU, WJP).

Utah. Washington: Leeds Canyon, G. F. Knowlton, W. J.
Hanson, T. H. Hsiao (1 9 , 3 3 , USU).

MEXICO

Lower California. 7 mi. sw La Paz, J. A. Chemsak (19,
CIS), 220 km s Tijuana, FXW (5 9 , CAS).

Tachysphex krombeiniellus sp.n.

ETYMOLOGY. — Named after K. V. Krombein
as a mark of friendship and gratitude.

DIAGNOSIS. — Tachysphex krombeiniellus is
similar to belfragei in having a bicolored gaster
and short thoracic vestiture which does not con-
ceal mesopleural sculpture. Unlike that species,
the propodeal dorsum of krombeiniellus is
evenly microareolate, and at least the apical
third of the hindfemur is red. It differs from
maurus and mundus by the shape of the clypeus
(female lip broadened, male middle section
slightly longer than wide). It can also be distin-
guished from marurus and most mundus by its
basally red gaster combined with the partly or
all red hindfemur.

GEOGRAPHIC DISTRIBUTION. — Mainly central
U.S. between northern Texas and North Da-
kota, west to 105th meridian, but also South
Carolina, Arkansas, and Florida.

MATERIAL EXAMINED. — HOLOTYPE: 9 , Florida, Levy Co.,
no specific locality, 3 June 1954, H. V. Weems (USNM).

PARATYPES: 22 9 , 14 3 ; June to Sep. Specimens for which
institution is not given below are all in UCD.

Arkansas. Mississippi: no specific locality, J. C. Nickerson

(1 9).

Colorado. Yuma: Yuma, collector unknown (1 3 ; 1 8,
WJP).

Florida. Gadsden: Quincy, Malaise trap (2 9 , UFG, WJP).
Levy: no specific locality, H. V. Weems (1 9 , CU; 2 d , FSCA;
1 9 , 1 cJ , USNM; 2 <S , WJP).

Kansas. Graham: Hill City, RRD (1 9). Pottawatomie:
Blackjack, H. E. & M. A. Evans, C. S. Lin, C. Yoshimoto
(2 d , MCZ).

Minnesota. Scott: Barden (between Savage & Shakopee),
C. E. Mickel (1 9, UMSP). Goodhue: Cannon Falls, C. E.
Mickel(l 9, UMSP).

Nebraska. Elaine: Halsey & Dunning, RRD (2 d). Box
Butte: Alliance, RRD (1 9). Dawson: Gothenburg, RRD (1
<S). Douglas: Omaha, collector unknown (19). Hall: 6 mi. w
Cairo, C. W. Rettenmeyer (1 <5). Lancaster: Lincoln, collector
unknown (1 9). Lincoln: North Platte, R. K. Schwab (1 9 ; 1
9 , WJP). Nance: Genoa, R. M. Barnes (19, INKS). Thomas:
Thedford, RRD (3 9 ; 1 9 , WJP).

North Dakota. Richland: 1 1 mi. w Walcott, J. R. Powers
(1 9, CIS).

South Carolina. Aiken: New Ellenton, A. Hook (1 9,
UGA).

Texas. Potter: 5 mi. N Amarillo, D. R. Miller (1 <S , USNM).

Wisconsin. Vernon: Genoa, collector unknown (2 9 ,
MPM).

Tachysphex menkei sp.n.

ETYMOLOGY. — Dedicated to A. S. Menke as
a mark of gratitude for his help.

DIAGNOSIS. — Tachysphex menkei can be eas-
ily recognized by the woolly setae of the head
and thorax; the finely, sparsely punctate scu-
tum, scutellum, mesopleuron, and propodeal
side; the largely impunctate, red gaster; and gla-
brous male sterna III-VI.

NATURAL HISTORY. — A female paratype is
pinned with her prey, a young nymph of a long-
horned decticine grasshopper, probably Ere-
mopedes sp., det. A. B. Gurney.

GEOGRAPHIC DISTRIBUTION. — Desert areas
from southwestern Texas to southern California.

MATERIAL EXAMINED. — HOLOTYPE: 9 , California, San
Diego Co., Borrego Valley, 19 Apr. 1957, R. M. Bohart
(UCD).

PARATYPES: 8 9 , 32 8 , Apr. to June.

Arizona. Graham: 18 mi. E Stafford, FGW & GDB (2 cJ,
UCD). Maricopa: 30 mi. E Gila Bend, R. F. Smith (1 <J , UCD).
Yavapai: 10 mi. NW Congress, FDP & LAS (1 9, UCD).
Yuma: 5 mi. SE Bouse, S. A. Gorodenski, JMD, MAC (1 6 ,
ASU).

California. Imperial: 30 mi. NE Glamis, R. R. Finger (2 <S ,
CSDA). San Bernardino: Adelanto, MEI (IS, UCD). San
Diego: Borrego Valley, H. R. Moffitt, EIS (2 S , UCD). Riv-
erside: Palm Springs, RMB (1 S, UCD); Andreas Canyon,
RMB, HKC (10 S , UCD, USNM, WJP).

42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 3

New Mexico. Otero: Alamogordo, collector unknown (1 LITERATURE ClTED

d, ANSP). Socorro: Bernardo, A. Apperson (1 9, USU).

., . , n... .. ... Xl(r™ D- D j BOHART, R. M., AND A. S. MENKE. 1976. Sphecid wasps of

Texas. Brewster: Alpine, J. Gillaspy (1 a, MCZ); Big Bend
... . T, , ,vr- ™ • . ™ x r> » i i- +, .1 • tne world. A generic revision. University of Ca hfornia

National Park (Nine Point Draw), R. Mason, J. F. McAlpme

/-, n -T » /-vr^ HUTO r>- n j XT .- i ™ i /o .1-1 Press, Berkeley, Los Angeles, London. 1 color pi., ix +

(3 9 , 7 <S ; CNC, WJP); Big Bend National Park (Santa Elena

Canyon), J. F. McAlpine (1 d , CNC); Big Bend National Park

«/ m D i AJ ,T j> n/-™ E-I n PuLAWSKi, W. J. 1971 . Les Tachysphex Kohl (Hym., Spheci-

(near Doughout Well), B. J. Adelson (2 <S , UCD). El Paso:

„ ... „ „ ii/TTO ,. j dae) de la region palearctique occidentale et centrale. Pan-

Tornillo, H. E. Evans & Rubink (1 9, WJP). Hudspeth: .

x* M j c c MO /-cm D_ -j- i stwowe Wydawmctwo Naukowe, Wrodaw. 464 p.

McNary H. E. Evans (19, CSU). Presidio: 3 mi. E Presidio,

n 9 Mr 7^ ' revision of the Neotropical Tachysphex

(Hym., Sphecidae). Polskie Pismo Ent. 44:3-80.
. 1977. A synopsis of Tachysphex Kohl (Hym., Spheci-

dae) of Australia and Oceania. Polskie Pismo Ent. 47:203-
332.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 4, pp. 43-58, 27 figs. November 4, 1982

A REVISION OF THE GRASSHOPPER GENERA

CHROMACRIS AND XESTOTRACHELUS
(ORTHOPTERA, ROMALEIDAE, ROMALEINAE)

By
H. Radclyffe Roberts

Department of Entomology, Academy of Natural Sciences,
19th and the Parkway, Philadelphia, Pennsylvania 19103

and
Carlos S. Carbonell*

Museu Nacional, Universidade Federal do Rio de Janeiro,
Quinta da Boavista, 20942 Rio de Janeiro, RJ, Brazil

ABSTRACT: Illustrations, diagnoses, and distributional records are given for 14 species and subspecies of the
Neotropical grasshopper genera Chromacris and Xestotrachelus, of which 1 species, Chromacris minuta, and 2
subspecies, C. trogon intermedia and C. psittacus pacificus, are described as new, and 4 names are newly
synonymized. Known information concerning food plants, imposition, habitat, and predator defense for Chro-
macris speciosa is briefly reviewed. (Orthoptera, Romaleidae, Romaleinae, grasshoppers, Neotropical, new taxa,
taxonomic revision)

INTRODUCTION treatment of the genus, a revisionary study ap-

The grasshopper species of the genus Chro- peared to be needed for this common and wide-

macris are of a striking color, usually a glossy spread group of grasshoppers. The monotypic

green with yellow markings, and red or yellow genus Xestotrachelus of southern South Amer-

wings. They occur in the humid areas of the ica is included in this study because of its sim-

American tropics from Mexico to Argentina. As ilarity to Chromacris and because it is the only

a general rule, but one species occurs at any one closely related genus.

locality. Adults are usually seasonal in their ap- The subfamily Romaleinae currently includes

pearance, so that some months of the year a about 48 genera. Rehn and Grant (1959) erected

species may appear to be absent from a local 16 tribes in this subfamily and proposed the tribe

fauna. Because most of the taxa of the genus Chromacrini for the genera Chromacris and

have been poorly defined or understood, the ap- Xestotrachelus. Because 10 of their tribes in-

plication of a number of their names has been elude but one or two genera each, and they give

uncertain, and there has been no comprehensive no distinguishing characters for these tribes, it

does not appear useful to recognize the tribe

* Boisista, Conselho Nacional de Desenvolvimento Cienti- Chromacrini and other such tribes of the

fico e Tecnologico, Brazil. subfamily.

[43]

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4

10 mm

Chromacris speciosa

FIGURE 1. Chromacris speciosa, Casupa, Florida, Uruguay.

The genus Chromacris includes at least eight
species, two of which are polytypic, each having
two subspecies, and two other species that we
tentatively recognize pending further informa-
tion. One species, C. minuta, and two subspe-
cies, C. trogon intermedia and C. psittacus pa-
cificus, are described as new. There are five
junior synonyms, three of which we newly syn-
onymize.

Specimens belonging to various collections
are indicated by the following abbreviations:
ANSP, Academy of Natural Sciences of Phila-
delphia; CACS, Dr. C. A. Campos Seabra Col-
lection; CSC, Carlos S. Carbonell Collection,
Montevideo, Uruguay; CHFR, C. H. F. Rowell
Collection, Zoologisches Institut der Universi-
tat, Basel, Switzerland; FCZ, F. Carrasco Col-
lection, Cuzco, Peru; MNHN, Museum Nation-
al d'Histoire Naturelle, Paris, France; MZSP,
Museu de Zoologia, Universidade Sao Paulo,
Brazil; UMMZ, University of Michigan Mu-
seum of Zoology, Ann Arbor, Michigan, U.S.A.

ACKNOWLEDGMENTS

We are grateful to our many friends for their
help and advice. We especially thank the follow-
ing persons and their respective institutions for
the loan of specimens. Dr. Irving J. Cantrall,
UMMZ; Drs. F. Carrasco and J. A. Escalante,
University of Cuzco, Peru; Dr. Marius Des-

camps and Christiane Amedegnato, MNHN; Dr.
Kurt K. Gunther, Museum fur Naturkunde,
Humboldt Universitat, Berlin, DDR; and the
late Dr. H. Reichardt, MZSP.

SPECIES LIST

In the following list of taxa we have attempted
to place similar or related taxa as close to one
another as practical. The number assigned to
each taxon matches the number in the text. Ju-
nior synonyms are given (in italics) below each
numbered taxon.

1 . Chromacris colorata (Serville)
Rhomalea pedes Pictet and Saussure

2. Chromacris minuta n.sp.

3. Chromacris miles (Drury)

4. Chromacris speciosa (Thunberg)

Acridium xanthopterum Hahn
Rhomalea stolli Pictet and Saussure

5. Chromacris nuptialis (Gerstaecker)

Rhomalea latipennis Pictet and Saussure

6. Chromacris trogon trogon (Gerstaecker)

7. Chromacris trogon intermedia n.subsp.

8. Chromacris psittacus psittacus (Gerstaecker)

9. Chromacris psittacus pacificus n.subsp.

10. Chromacris icterus (Pictet and Saussure)

Rhomalea opulenta Gerstaecker

1 1 . Chromacris peruviana (Pictet and Saussure)

12. Xestotrachelus robustus (Bruner)

Xestotrachelus hasemani Bruner

ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS

Xestotrachelus robustus

FIGURE 2. Xestotrachelus robustus, Chapada dos Guimaraes, Mato Grosso, Brazil.

KEY TO SPECIES OF CHROMACRIS 5(4').

1. Exterior of small aedeagus membra-
nous (Figs. 19-20). First two plaits or
anterior fold of wings entirely black to
base (Figs. 3-6). Colorata group ____ 2

1 ' . Exterior of aedeagus sclerotized form-
ing a short collarlike structure around * •
the phallotreme opening (Figs. 22-25).
First two plaits or anterior fold of
wings yellow on basal half (Figs. 7-9),
or some black on basal half of first
plait (Fig. 10). Trogon group 6

2(1). Yellow on part of antennae. Stridulat-
ing structures weakly developed (Fig.
15)

2'. Antennae entirely black. Stridulating
structures well developed (Figs. 16-
17) .

3(2). Basal portion of antennae yellow
(Mexico to Costa Rica) C. colorata

3'. Distal portion of antennae yellow
(Acapulco, Mexico) C. minuta

4(1')- Membrane of tegmina dark brown to 9(7').

black with strongly contrasting straw-
colored veins (inland south central
Brazil) C. nuptialis

4'. Not as above _ 59'.

6(1').
6'.

7(6).

7'.

8(7).

Posterior yellow margin of pronotum
interrupted by black or green at angle
between disc and lateral lobes. Rim of
coxal articulation on mesathorax and
metathorax entirely or partially yellow

(South America) C. speciosa

Posterior yellow margin of pronotum
not interrupted at angle between disc
and lateral lobes. Rim of coxal artic-
ulation on mesathorax and metathorax
entirely green (southeastern Mexico)

C. miles

Antennae entirely black _.
Antennae with yellow tips

7
10

No bands on hind tibiae 8

Yellow bands on hind tibiae 9

Yellow bands lacking on all legs (Cos-
ta Rica) C. trogon trogon

Yellow bands on hind femora (Guate-
mala, Belize, and Honduras)

C. trogon intermedia

Proximal yellow band on hind femora
interrupted on outer ventral portion
(Costa Rica to northern Colombia and

Venezuela) C. psittacus psittacus

Proximal yellow band on hind femora

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4

X. robustus

FIGURES 3-11. Male hind wings of seven of the eight species of Chromacris (3-10) C. nuptialis being omitted because of
similarity to C. speciosa (6), and Xestotrachelus robustus (11), all at same scale and from the following localities: (3) colorata
Medellin de Bravo, Veracruz, Mexico; (4) minuta Acapulco, Guerrero, Mexico; (5) miles Boloyuc, Quintana Roo, Mexico;
(6) speciosa Aratinga, Rio Grande do Sul, Brazil; (7) trogon San Lorenzo, Alajuela, Costa Rica; (8) psittacus Cabima, Panama;
(9) icterus Tabatinga, Amazonas, Brazil; (10) peruviana Satipo, Junfn, Peru; (1 1) A', robustus Cerro Cora, Amambay, Paraguay.

ROBERTS AND CARBONELL: REVISION OF CHROMACR1S AND XESTOTRACHELUS

entire (western Colombia and western
Ecuador) C. psittacus pacificus

10(6'). Hind tibiae with single, distal, yellow
band (southeastern Colombia to north-
eastern Peru) C. icterus

10. Hind tibiae with two yellow bands
(Peru) C. peruviana

Chromacris Walker

Chromacris WALKER, 1870:643. [Type-species Gryllus spe-
ciosus Thunberg, 1824, by subsequent designation of Rehn
1904:532.]

DIAGNOSIS. — Neither pronotal crest nor pro-
truding fastigium present; fully alate (Fig. 1).
Medium size, body length of females 33 mm (C.
minutd) to 55 mm (C. icterus and C. peruviana).
Glossy olive-green to dark green with contrast-
ing yellow or yellow tinged with red markings.
These markings may be much reduced as in Pe-
ruvian specimens of C. speciosa. Hind wings
various shades of red, orange, or yellow with
contrasting black pattern (Figs. 3-11) character-
istic of this genus and Xestotrachelus, distin-
guishing them from all other genera of subfam-
ily. Genitalia (Figs. 19-26) rather uniform
throughout the genus. Noteworthy are weakly
developed aedeagal valves.

CLASSIFICATION. — Two groups of Chroma-
cris can be recognized. One group including tro-
gon, psittacus, icterus, and peruviana (trogon
group) have the proximal half of the first two
plaits of the hind wings without black on the
anterior portion of first two plaits (Fig. 10). In
contrast, first two plaits of group containing co-
lorata, minuta, miles, speciosa, and nuptialis
(colorata group) are entirely black (Figs. 3-6).
Aedeagal valves of trogon group are externally
sclerotized (Fig. 22-25) whereas those of the co-
lorata group are small membranous lobes
(Figs. 19, 20). Trogon group members common-
ly have yellow wings, occasionally orange,
and rarely red. Wings of colorata group mem-
bers are commonly red, orange in some geo-
graphical areas, and frequently yellow in part
of Atlantic coastal area of Brazil. In the trogon
group, prosternal tubercle small and pointed
with concave sides as seen in profile. In con-
trast, C. miles, C. speciosa, and C. nuptialis
have a large, bluntly pointed tubercle that is
variable in form, even intra-specifically. On the
other hand, tubercle of C. colorata and C. mi-
nuta more nearly resembles that of the trogon

group in its small size, may be pointed, but sides
in profile are rarely concave. C. miles and its
two closely related species, C. speciosa and C.
nuptialis, have a well-developed stridulatory ap-
paratus, as in many other species of the Ro-
maleinae. The serrate cross veins (SV) between
the first (1A) and second (2 A) anal veins of the
second plait (Figs. 16, 17) contact the raised
scraper veins on underside of tegmen when
wings are folded. Tympanate or fenestrate areas
(TA) on either side of these cross veins are well
developed as resonators. This stridulatory ap-
paratus is more weakly developed in other
species of Chromacris and, in many cases, may
not be functional. For example, compare the
wing of C. icterus (Fig. 15) with those in Figures
16 and 17. In summary, the trogon group is dis-
tinguished from the colorata group by the ae-
deagal valves and black pattern of hind wings,
and the colorata group is divided into two
subgroups by shape of prosternal tubercle and
stridulatory apparatus.

Recognition of species in the genus Chroma-
cris has been difficult because of the lack of
morphological characters and dependence on
color and color pattern. The trogon group of four
species, including two subspecies, can be rea-
sonably well defined by a combination of differ-
ent color-pattern characters (Table 2). The illus-
trations of the aedeagus of this group (Figs. 22-
25) may suggest species differences, but individ-
ual variation is such that clear distinctions be-
tween species are not evident. The wide-ranging
C. speciosa of South America has been most
puzzling because of the great amount of geo-
graphical variation of color and color pattern
and variation within a local population. It has
been difficult to decide whether we are dealing
with species, subspecies, or just color forms. As
an example of color forms, about half of the 21
specimens of C. speciosa recorded from Flores-
ta de Tijuca near Rio de Janeiro, Brazil, have
yellow wings and the other half orange wings,
with no intermediates. We have concluded that
it is most practical to treat C. speciosa, at least
for the present, as but one highly variable
species. We need more information on C. miles
and C. nuptialis to understand their status as
species and their relationship to C. speciosa.

GENERAL OBSERVATIONS. — Most of the avail-
able data for the genus refer to C. speciosa. The
biology of this species was studied in Tucuman,

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4

,2Aa

FIGURES 12-14. Male hind wings of three species of Chromacris showing relative position of radius (R) median (M) fork to
the second anal accessory (2Aa) fork. The 2Aa fork is more distant from the base of wing than the R-M fork in C. speciosa
(Fig. 14) and its related species, C. miles and C. nuptialis, whereas in C. trogon and C. psittacus (Figs. 12 and 13) and all other
species of the genus, the 2Aa fork is equidistant or closer to base of wing than the R-M fork. The If is the first and the 2f is
the second fold line of wing.

FIGURES 15-18. Details of stridulatory area of male hind wings of three species of Chromacris (Figs. 15-17) and Xesto-
trachelus robustus (Fig. 18). Note well-developed tympanate areas (TA) in Figs. 16-18. Terminology of wing venation follows
Ragge (1955). SV, serrate veinlets or cross veins; TA, tympanate or fenestrate areas; R, radial vein; Rs, radial sector; Cul,
first cubital vein; 1A, first anal vein; 2A, second anal vein.

Argentina, by Barrera and Turk (1977). Data on
the biology of C. colorata have been reported
from Monterrey, Mexico, by Pretto-Malca
(1968), at the other extreme for the distribution

of the genus. Some data on the food of other
species exist, mainly in papers of applied ento-
mology. Some aspects of the general biology of
the species of the genus, such as their general

ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS

preference for solanaceous and composite plants
and the gregariousness of their juveniles, are
generally known by all entomologists who have
collected these insects in the field.

OVIPOSITION AND DEVELOPMENT. — EggS of

C. speciosa are laid in the soil, the top of the
egg-pod 10 to 20 mm under the surface. Eggs
are cemented together in the pods, but not
embedded in the frothy secretion which covers
the egg-pods of other acridoids. This secretion
just forms the upper half of the pod, while the
egg-mass is bare and usually 15 mm long, 8.5
mm wide. The number of eggs in each pod varies
between 61 and 70, with a mean of 66 eggs (Bar-
rera and Turk 1977). This species grows from
hatching to imago, under the Tucuman climate,
in 30 to 60 days, passing through five instars in
the male and six instars in the female. Nymphs
are black with red and some white markings.
Sexual maturity is attained 10 days after the last
molt. Adults mate repeatedly and females lay at
least two pods. The insects usually disperse af-
ter reaching the imaginal stage (Turk and Bar-
rera 1976). Pretto-Malca (1968) stated that egg-
pods of C. colorata contain an average of 35
eggs and that the insect reaches the imaginal
stage through six nymphal instars, and sexual
maturity about 18 days later. Nymphs of this
species are highly gregarious.

FOOD PLANTS. — Chromacris speciosa prefers
solanaceous plants. Turk and Barrera (1976) re-
ported its feeding on Cestrum parqui, C. stri-
gillatum, C. lorentziana, Lycium cestroides,
Solarium argentinum and 5. verbascifolium (So-
lanaceae), and also on Verbesina encelioides
(Compositae) and alfalfa (Medicago saliva, Le-
guminosae). They bred to maturity nymphs of
this species found on alfalfa, using only this
plant for food, and development was normal.
They reported that in laboratory breedings
nymphs which were first fed on Lycium ces-
troides readily changed to other species of Lyci-
um, but would starve to death rather than accept
Solanum or genera of other plants. Conversely,
nymphs started on Solanum would not accept
Lycium. Thus, food plants, at the generic level,
are determined by the first food of the newly
hatched nymphs. The same feeding experiments
demonstrated that this species would not eat
species of the grass family (Turk and Barrera
1976). Carrasco (1962), however, reported C.
speciosa and another unidentified species (evi-

dently C. peruviana according to Carrasco' s de-
scription) feeding on rice plants and doing con-
siderable damage to this crop in Peru. Guagliumi
(1973) mentioned C. speciosa as feeding on sug-
ar cane in northeastern Brazil. Astacio-Cabrera
(1975) reported C. colorata in Nicaragua on the
composite Baltimora recta, and Pretto-Malca
(1968) has stated that this species in Mexico
usually feeds, and has been bred on, Solanum
elaeagnifolium. Rowell (1978) reported the So-
lanaceae as the preferred food of C. trogon in
Costa Rica.

BEHAVIOR. — The gregarious stages of Chro-
macris, which are usually seen forming large
groups on the tops of their food plants, together
with their bright and contrasting coloration sug-
gest that they are unpalatable or poisonous to
predators and that their coloration is premoni-
tory. One of us (H.R.R.) recently observed 20
to 30 conspicuous nymphs on top of a tussock
of grass two or three meters from a small sola-
naceous shrub stripped of its leaves, which sug-
gests that the gregarious behavior is a part of
the premonitory defense. After reaching the
imaginal stage, these insects tend to disperse.
Adults are very visible during flight, but once
they alight on vegetation, they seem to disap-
pear after the display of their colorful wings sud-
denly ceases.

HABITAT. — The species of this genus are usu-
ally found at forest edges and clearings, road-
sides, edges of cultivated fields, and nearly all
places where herbaceous solanaceous and com-
posite plants occur. They seem to avoid heavy
forest and prairie habitats.

1 . Chromacris colorata (Serville)

Acridium coloratum SERVILLE, 1839:674. [Holotype, MNHN,
no longer extant, said to come from South Carolina, USA,
obviously in error. Mention of the 8-10 basal segments of
the antennae as yellow and others black clearly indicates
that it is the Mexican species.]

Rhomalea pedes SAUSSURE, 1859:392. [Lectotype, (J.Geneva
Museum; Mexico. So labelled by C.S.C. and here desig-
nated.]

DIAGNOSIS. — Eight to 12 proximal segments
of antennae yellow, distal segments black — a
unique color pattern for the genus. Posterior
margin of the pronotum yellow, but no yellow
patches on midportion of lateral lobes, as is usu-
al in C. speciosa. Three yellow bands on hind
femur and only one distal band on hind tibia.
Middle leg with one band on tibia, femur, and

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4

22 C. trogon

23 C. psittacus

C. icterus

FIGURES 19-25. Dorsal and lateral views of aedeagi. Figs. 19 and 20 are examples of colorata-miles group. Figs. 22-25 are
examples of trogon group. Fig. 21. Xestotrachelus robustus from Cerro Cora, Amambay, Paraguay. Fig. 19. Chromacris
colorata from Medellin de Bravo, Veracruz, Mexico. Fig. 20. C. speciosa from Resistencia, Chaco, Argentina. Fig. 22. C.
trogon from San Lorenzo, Aulajuela, Costa Rica. Fig. 23. C. psittacus from Las Pavas, Santa Marta Mts., Colombia. Fig. 24.
C. icterus from Villavicencio, Colombia. Fig. 25. C. peruviana from Aucayacu, Huanuco, Peru.

trochanter. Fore legs with one band on tibia.
Hind wings rich cherry-red, with black pattern
(Fig. 3) similar to that in C. speciosa. Prosternal
tubercle tapers to a narrow point, much as in C.
psittacus and other yellow- winged species. The
small, short aedeagus formed by a pair of papil-
lose, flattened lobes (Fig. 19) slightly sclerotized
internally. Aedeagus similar to others of the co-
lorata group.

DISTRIBUTION. — Tropical humid areas of
Mexico south to Costa Rica. Of common occur-
rence July to September.

SPECIMENS. — Mexico. States of Nuevo Leon, Tamaulipas,
San Luis Potosi, Veracruz, Oaxaca, Guerrero, Morelos, Ja-
lisco, Nayarit, Sinaloa, and Yucatan.

Guatemala. EL PETEN: 3 mi [4.8 km] S Tikal, 16 Aug. 1974

(J. C. Lee), 6d, 39. 22 mi. [35 km] NW Poptun, 15 Jul. 1974
(J. C. Lee), IcJ, 39.

Belize. Rio Grande, Aug. 1931 (J. J. White), 39.

Nicaragua. MANAGUA: Sep. 1955 (E. Morales- Agacino),
2<J, 19, CSC.

Costa Rica. GUANACASTE: Canas, Sep. 1965 (C. H. F. Row-
ell), 1 cJ , 1 9 , CHFR.

COMMENTS. — Specimens from Volcan Colima
(Jalisco, Mexico) and Guanacaste (Costa Rica)
agree with Serville's description of Acridium
coloratum in lacking any banding on fore and
middle legs and on hind tibia, and much reduced
banding on hind femora. The type-specimen of
Rhomalea pedes has fully banded legs, as have
most Mexican specimens. Costa Rican speci-
mens show small yellow marks on the prozonal
part of the lateral lobes of the pronotum, as is

ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS

also true of some specimens of C. speciosa.
These Costa Rican specimens also have 12 seg-
ments of the antennal flagellum yellow, the fol-
lowing 4 segments part black and part yellow,
and only the tip of the antennae entirely black.
The Costa Rican specimen has no yellow on
posterior margin of pronotum.

2. Chromacris mi nut a n.sp.

DIAGNOSIS. — Six to eight apical antennal seg-
ments yellow, other segments black. Hind wings
orange-red rather than cherry-red as in C. co-
lor ata. Body color pale olive-green with greatly
reduced yellow markings. No yellow bands on
legs except weakly developed proximal and me-
dian bands on hind femur. Prosternal tubercle
short and conical. End of aedeagus much as in
C. colorata. Body size small for the genus,
males 26-31 mm, females 35-39 mm from fas-
tigium to end of wings.

SPECIMENS. — Holotype: cJ, ANSP, Acapulco, Guerrero,
Mexico, 13 Aug. 1935 (H. R. Roberts, E. R. Helwig).

Paratypes: Mexico. GUERRERO: 5 mi. [8 km] N Acapulco, 15
Sep. 1940 (C. Bolivar, H. R. Roberts), 7cJ , 19. Same data as
for holotype, 63 , 8 9 , 3 juv.

COMMENTS. — This species is most similar to
C. colorata. It should be looked for elsewhere
along the Pacific coast of Mexico. A female of
C. colorata from between Tierra Colorada and
Rio Papagayo, about 40 km north of Acapulco,
shows some reduction of yellow banding of the
legs as occurs in C. minuta, but in other respects
is typical of C. colorata.

3. Chromacris miles (Drury)

Gryllus locusta miles DRURY, 1773:79, pi. 42, fig. 2. [Holo-
type, 9 , not found in British Museum or Oxford collections,
but excellent figure should suffice; "Bay of Honduras."]

DIAGNOSIS. — Red wing pattern (Fig. 5) similar
to that in C. colorata and others of the colorata
group. Resembles C. colorata in lacking yellow
in middle area of lateral lobe of pronotum. This
condition occurs rarely in C. speciosa. Similar
to C. speciosa in having entirely black antennae
and two yellow bands rather than one on hind
tibiae, but differs in having yellow on hind mar-
gin of pronotum interrupted by black or green
only on midline, whereas in C. speciosa it is
interrupted on midline and both sides at the an-
gles that limit disc from lateral lobes of meta-
zona. Also, in C. miles rim of coxal articulation

on mesathorax and metathorax green, whereas
it is entirely or partially yellow in C. speciosa.

SPECIMENS. — Mexico. QUINTANA Roo: 5 mi. [8 km] SE Po-
lyuc (Boloyuc), 28 Jul. 1960 (P. M. Litchfield), IcJ, UMMZ.
VERACRUZ: Laguna Verde, Aug. 1974 (M. Descamps), a small
series of males and females, MNHN.

COMMENTS. — It has been thought that Dru-
ry's name should be applied to Thunberg's
South American species, C. speciosa. Finding
specimens from Veracruz and the peninsula of
Yucatan which closely match Drury 's figure
supports the existence of a distinct species in
the Bay of Honduras region. Based on the black
pattern of the hind wings, this species belongs
to the colorata group and is closest to C. spe-
ciosa in the strong development of the stridu-
latory areas on the hind wing. Its color pattern
is also more similar. C. speciosa, however, does
not occur north of Colombia, and C. miles oc-
curs within the range of C. colorata. More in-
formation on the distribution of the genus in this
region is needed to clarify our understanding of
Drury's species.

4. Chromacris speciosa (Thunberg)

Gryllus spedosus THUNBERG, 1824:404. [Lectotype, 9 , so la-
belled by C.S.C. and here designated; two male syntypes
also examined; Uppsala Museum; Brazil].

Acridium xanthopterum HAHN, 1835, table A, fig. 2. [Holo-
type unknown; Brazil. Hahn attributes the name to Perty in
"Ins. nov bras.," but no such reference has been found.
Black pattern of hind wing shown in illustration identifies it
as the yellow-winged form of this species. New synonym.]

Rhomalea stolli PICTET AND SAUSSURE, 1887:351. [Lecto-
type, cJ, so labelled by C.S.C. and here designated; Geneva
Museum; Bahia, Brazil. Synonym by Kirby 1910:373.]

DIAGNOSIS. — Varies geographically and local-
ly. Red winged over most of its range, but in
lowland coastal area from Bahia, Brazil, to Uru-
guay, wings are frequently orange or yellow, and
tegmina tend to be green rather than green
tinged with red. Yellow-winged individuals
readily distinguished from the typically yellow-
winged species of the trogon group by entirely
black basal half of the first two plaits (anterior
or first paired fold of wing) (Fig. 6). In Paraguay,
Argentina, and Uruguay wings tend to be or-
ange-red, and yellow markings are tinged with
red. In Bolivia, Peru, and Ecuador reduction
and variation in leg banding evident. Elsewhere,
hind femur almost always has three pale bands
and the hind tibia two pale bands. Specimens
from Ecuador and Peru lack hind tibial bands,

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4

TABLE 1. VARIATION IN BANDING OF HIND FEMUR OF
Chromacris speciosa (22 specimens from Ecuador and Peru,
60 specimens from Santa Cruz, Bolivia).

Banding of hind femur

Ecuador, Peru

Bolivia

Strong Weak Absent Strong Weak Absent

Proximal

—

Median

—

Distal

—

and hind femur usually has a strong proximal
band, median band may be weak or absent, and
distal band nearly always absent. Some 60 spec-
imens from city of Santa Cruz region of eastern
Bolivia show great variation in presence or ab-
sence of various bands. Hind tibia may have a
distal band or none. Hind femur usually has
strong proximal band; it is never absent; median
band usually weakly developed; distal band may
be strongly developed (Table 1). Also, in the
Andean region north to Ecuador, body color and
tegmina darker.

SPECIMENS. — Unless otherwise noted all specimens have
red hind wings, three yellow bands on hind femur, two yellow
bands on hind tibia, and tegmen tinged with red.

Colombia. MAGDALENA: Aracataca, 4-10 Aug. 1920 (Rehn,
Hebard), 8d, 39; (3 lack proximal tibial band).

Venezuela. CARABOBO: San Esteban, Nov.-Dec. 1939 (P.
Van Duse), 6d, 79. COJEDES: 26 km s jet. Rts. 8 and 13 on
Rt. 8, forest, 13 Jul. 1981 (Otte et al.), Id, 39 .

Guyana. Bartica, Dec. 1912 (H. S. Parish), 30d, 359.

Brazil. AMAPA: Rio Puxaca, Mazagao, Feb. 1961 (J. C. M.
Carvalho), 39, UMMZ. Eighteen specimens from Belem,
Santarem, Obidos, and Manaus on the Amazon are similar to
the Guyana series. PARA: Jacareacanga, 6°16'S, 57°44'W,
Dec. 1968 (Alvarenga), 3d, 99 , UMMZ. BAHIA: 100 km NW
Feira de Santana, 13 Mar. 1981 (Roppa, Carbonell, Roberts),
2d , 29 ; Itabuna, research center, 22 Nov. 1974, cacao forest
(Roberts, Carbonell), 19; Mucuri, Aug. 1977 (Roppa, Beck-
er), 2d (one yellow wings, other orange wings). ESPIRITO
SANTO: Itapemirim falls, 1-6 km E BR. 101, edge of forest and
marsh, 5 Dec. 1974 (Roberts, Carbonell), 49 (lack usual yel-
low median spots on lateral lobe of pronotum); 3 km s Lin-
hares, cacao forest, 1 Dec. 1974 (Roberts, Carbonell), 2d, 19
(lack usual yellow spots on lateral lobe of pronotum). Rio DE
JANEIRO: Floresta de Tijuca, Jan. 1981, 4d, 69 (yellow
wings), 5d, 69 (orange wings); BR. 101, 1 Feb. 1974 (D.
Otte), 26 (yellow wings, tegmina lack red tinge); Petropolis,
12 Apr. 1913 (M. Burr), 2$ (yellow wings, tegmina lack red
tinge). MINAS GERAIS: Vigosa, 9 Aug. 1938 (B. T. Snipez),
29 (yellow wings, tegmina lack yellow tinge); 46 km SE Ita-
juba, 1400 m, 21 Mar. 1980 (Roppa, Carbonell, Roberts), 3d,
39. GOIAS: betw. Sao Simao and Jatai, 5 Mar. 1980 (Roppa,
Carbonell, Roberts), Id, 29; 15-30 km E Mineiros, 7-9 Mar.
1980 (Roppa, Carbonell, Roberts), 1 9 . SAO PAULO: 10 km w
Sao Joao de Boa Vista, 19 Mar. 1980 (Roppa, Carbonell, Rob-
erts), 30d; Franca, Jan. 1911 (E. Garbe), Id, 19; Salto

Grande, Feb. 1911 (H. Luderwaldt), Id; Cubatao (Alin), 2d,
19 (orange wings, tegmina lack red tinge); Piracicaba, Id
(orange wings). PARANA: 24°38'S, 54°07'W, 500 m, Mar. 1965,
"virgin deciduous forest (no Araucaria) with many palms
(Euterpe etc.), no grass, under growth of ferns and other
plants" (F. Plaumann), 4°, 199, UMMZ; Curitiba, 13 Feb.
1941 (J. R. Bailey), Id, 19, UMMZ (male has orange wings,
lacks red tinge on tegmina). SANTA CATARINA: Nova Teuton-
ia, 27°H'S, 52°23'W, 6 km sw Seara, 300-500 m, 1961-1964
(F. Plaumann), 6d, 59, UMMZ; Corupa, Jan.-Mar. 1956-
1 962 (Anton Mailer) , 3 d , 9 9 , UMMZ ( 1 d , 5 9 have red wings ,
2d lack red tinge on tegmina, 2d, 49 have orange-yellow
wings and lack red tinge on tegmina); Rio Capivari, 1889
(Fruhstorfer), 29 [gift of Dr. H. Saussure, ANSP, labelled R.
miles Drury and Rhomalea speciosa, probably what Pictet
and Saussure considered to be R. miles Var. C, as it has
yellow wings; there are two Rio Capivari 's in«astem lowlands
of Santa Catarina]; Pinhal 700 m, Apr. 1959 (Anton Mailer),
1 9 (yellow wings, lacks red tinge on tegmina). Rio GRANDE
DO SUL: Aratinga, Feb. 1964 (Carbonell, Mesa, Monne), Id
(yellow wings). MATO GROSSO: 40 km E Rodonopolis, 1 1 Mar.
1980 (Roppa, Carbonell, Roberts) Id; 30 km NW Alto Ara-
guaya, 750 m, 10 Mar. 1980 (Roppa, Carbonell, Roberts) Id;
Chapada near Cuiaba, 4d, 39 (Id lacks distal band on hind
femur); Corumba, Urucum, 22-29 Dec. 1919 (R. G. Harris),
12d, 129 (proximal median bands weak, distal band strong
on hind femur, proximal tibial band usually absent); Tres La-
gos, 6-10 Dec. 1919 (Harris), 49 (hind legs fully banded, and
one of these with yellow markings strongly tinged with red),
1 9 (distal band on hind femur and hind tibia only). MATO
GROSSO SUL: 30-60 km E Aquidauana, 16 Mar. 1981 (Roppa,
Carbonell, Roberts), 26.

Uruguay. Whole country, Dec.-Apr., large series, CSC
(yellow markings tinged with red).

Ecuador. Balzapamba (R. Haensch), Id (weak proximal
band on hind femur). Putumayo Dist., La Sombra to El En-
canto, 23 Aug. 1920, Id (strong proximal and weak median
band on hind femur).

Peru. JUNIN: Satipo, 15 Nov. 1945 (P. Paprzycki), Id, 89;
Satipo, 650 m, Jul. 1940 (Schunke) 1 9 ; Col. Perene, El Cam-
pamento, 22 Jul. 1920, 29 ; Chanchamayo, Id, 19; Vilcanota,
1 9 ; Puerto Yessup, Feb. 1930 (M. A. Carriker), 1 9 . Cuzco:
Valle de Urubamba, Sahayaco, 800 m, 7 Dec. 1947 (Wey-
rauch), 4d; Prov. La Convention, Sangobatea, Jan. 1976 (J.
S. Escalante), 3d, 39, JAE; Prov. Paucartambo, Salvation,
Oct. 1968 (F. Carrasco), 1 9 , FCZ. All Peruvian specimens
have moderate to well-developed proximal bands, weak to no
median bands, and no distal bands on hind femur; no bands
on hind tibia and other legs.

Bolivia. SANTA CRUZ: Province of Sara, 450 m, Jan. 1918
(J. Steinbach), 19d, 279 (35 had no tibial banding, 11 had
weak distal yellow bands; on hind femur all had weak to strong
proximal bands, 14 had no median bands, 22 had weak to
strong distal bands, and 24 had no distal bands) [Note: this
previously recognized Province of Sara, bounded in part by
the Rio Grande or Guapay and the Rio Mamore or Ichilo, is
the region where Steinbach did much of his collecting, and
included the town of Buena Vista (where his relatives lived
and where a niece presently operates a small restaurant, store,
and inn), Portachuelo, and the city of Santa Cruz, that is,
Santa Cruz de la Sierra.]; Buena Vista, May 1917 (Steinbach),
2d, 19; between Buena Vista and San Carlos, 350 m, cacao
forest, 21 Feb. 1976 (Ronderos, Roberts) 2d, 19; between
Buena Vista and Portachuelo, 20 Feb. 1976, 19 (specimens
from last three localities similar in variation to those from

ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS

Prov. of Sara; Santa Cruz de la Sierra, Feb. 1922 (J. Stein-
bach), 5<J, 69; 18 km sw Santa Cruz, 400 m, 16 Feb. 1976
(Ronderos, Roberts), 36, 29 (of last 16 specimens, 15 had a
strong distal band on hind femur, 16 lacked this band); Prov.
of Nuflo de Chavez, Ascencion, 15°42'S, 63°05'W, 500 m,
Nov. 1963 (Walz), 46, 59 (5 had distal and 4 had no distal
band on hind femur). See Table 1 for a summary of this band-
ing.

Paraguay. 26 specimens from Villa Rica, Jan., Feb.; Sapu-
cay, Jan.-Apr.; Horqueta, Dec., Jan. (yellow markings
strongly tinged with red; hind femur with three bands, hind
tibia with two bands).

Argentina. 102 specimens from 24 localities in the provinces
of Jujuy, Feb.; Salta, Mar.; Chaco, Feb.; Misiones, Dec.,
Feb.; Tucuman, Mar.; Catamarca, Mar.; Cordoba, Mar.;
Entre Rios, Mar.; La Rioja, Feb.; Mendoza Feb.-Apr.; San
Luis, Jan.; Buenos Aires, Feb. As in Paraguay and Uruguay,
yellow markings are strongly tinged with red; hind femur with
three bands, hind tibia two bands. Adults may be found De-
cember to April.

COMMENTS. — Unlike other species of the ge-
nus, no consistent differences have been found
to distinguish various geographical develop-
ments. Comparing specimens, for example,
from Carabobo, Venezuela, with those from
Santa Cruz, Bolivia, or Corupa, Brazil, it is ev-
ident that considerable geographic differentia-
tion occurs, but it does not seem possible or
practical with our present evidence to recognize
subspecific elements of the species.

5. Chromacris nuptialis (Gerstaecker)

Rhomalea nuptialis GERSTAECKER, 1873:185. [Holotype, 6,
bearing label with number 2008, and 9 allotype, Berlin Mu-
seum; Saltogrande (Sellow). The locality of Salto Grande of
Sellow is uncertain. Sellow visited Salto Grande on the
Uruguay River (Department of Salto, Uruguay), but none
of the species he labelled Saltogrande has ever been found
in Uruguay or the adjacent Brazilian state of Rio Grande do
Sul. Sellow's Saltogrande is very probably that on the Par-
anapanema River in the state of Sao Paulo. Types exam-
ined.]

Rhomalea latipennis PICTET AND SAUSSURE, 1887:351. [Holo-
type, 6 , Geneva Museum; Brazil. It does not have a locality
label but bears the number 477-56, which in the museum
records corresponds to Brazil, collected by Ferrier circa
1856. We have been unable to trace the collector's route in
Brazil. Holotype examined. New synonym.]

DIAGNOSIS. — Coloration highly variable, in-
cluding individuals with pale yellow and pale red
wings. Body and legs with yellow or red mark-
ings (irrespective of wing color). Antennae black.
Pattern of hind wings as in C. speciosa. Tegmina
very characteristic, membrane dark brown to
black, strongly contrasting straw-colored veins.
Fore and middle legs without transverse bands,
ground color variable from greenish yellow to
reddish brown; longitudinal series of black spots,
sometimes coalescing into black streaks. Hind

femora black to dark brown, with longitudinal
carinae and fishbone pattern of a lighter color,
variable from reddish brown to straw-yellow;
transverse bands absent or very slightly marked,
only exceptionally, plainly visible, proximal one
on upper half only, median one may be complete,
distal one always absent. Proximal and median
bands always visible on inner and lower surfaces
of hind femur as conspicuous yellow or red areas,
no trace of distal one. Hind tibia generally dark
colored, especially on inner side, transverse
bands generally absent, sometimes faintly
marked, the distal one, proximal one, or both
may be visible in different specimens. Prosternal
tubercle relatively slender, long, and curved rear-
wards.

DISTRIBUTION. — Inland south-central Brazil,
including southern Goias, western Minas Gerais,
western Sao Paulo, and northwestern Parana.

SPECIMENS.— Brazil. D.F.: Brasilia, Nov. 1963 (N. Tanger-
ini), 26 (yellow wings), CACS. GOIAS: rodovia Anapolis-
Brasilia, km 63, 17 Feb. 1964 (H. M. Canter), 16 (yellow
wings), MZSP; betw. Sao Simao and Jatai, 5 Mar. 1980
(Roppa, Carbonell, Roberts), 19 (red wings), CACS; Min-
ieros, Feb. 1975 (Roppa, Silva), 19 (red wings), CACS.
MINAS GERAIS: Uberaba, Feb. 1979 (Roppa, Silva), 16 (red
wings), CACS; Diamantina, Mar. 1956 (D. Albuquerque), 1 9
(red wings), CACS; Lagoa Santa, Jul. 1965 (M. S. Morgante),
16 (yellow wings), MZSP. PARANA: Vila Velha, Jan. 1975
(C. Valle), 16, 19 (yellow wings), MZSP.

COMMENTS. — Individuals of this species are
highly variable in color, and therefore difficult
to identify. However, the only other species
known from the area is C. speciosa, from which
it can be separated by the peculiar coloration of
its tegmina, the very different color on pronotum
and legs, and the form of its pronotal tubercle.
The species appears to be uncommon, being al-
ways represented by one or two specimens from
each locality, which is unusual for species of this
genus. Possibly, this taxon is a highly aberrant
variation of C. speciosa in the middle of whose
territory it occurs, but the constancy of some of
its characters seems to indicate that it is a dis-
tinct species.

6. Chromacris trogon trogon (Gerstaecker)

Rhomalea trogon GERSTAECKER, 1873:186. [Holotype, 19,
Berlin Museum; Costa Rica. (Gerstaecker noted that hind
legs were lacking. A bright yellow marked leg was later
attached and now removed. Holotype examined.]

DIAGNOSIS. — Hind wings yellow to orange-
yellow, and lack black on anterior basal half of
first two plaits (Fig. 7). Antennae entirely black.
No yellow banding on legs, although often faint
pale green bands present on hind femora (Table

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4

TABLE 2. DIAGNOSTIC CHARACTERS FOR THE SPECIES AND SUBSPECIES OF THE TROGON GROUP OF GENUS Chromacris. A
band that does not extend entirely around the hind femur is listed as a half band.

Name

Range

Antenna

Basal Vi
of wing

Pronotal
metazona

Hind femur

Hind tibia

/. intermedia

N Guatemala
to Honduras

all black

all yellow

no yellow

3 half bands

no bands

t. trogon
p. psittacus

Costa Rica
Costa Rica to
N Colombia,
N Venezuela

all black
all black

all yellow
all yellow

no yellow
no yellow

no bands
2 and Vi bands

no bands
2 bands

p. pacificus

w Colombia,
w Ecuador

all black

all yellow

no yellow

3 bands

2 bands

icterus

E Colombia
to NE Peru,
NW Brazil

tip yellow

all yellow

yellow patches

3 bands

1 band

peruviana

Peru

tip yellow

plait 1 black

yellow patches

3 bands

2 bands

2). Reproductive structures and prosternal tuber-
cle similar to those in C. psittacus and other
yellow-winged species of the trogon group.

DISTRIBUTION. — Costa Rica and questionably
Nicaragua.

SPECIMENS. — Costa Rica. Pozo Azul, Rio Ferris or Parrita
(forested foothills of Pacific coastal plain), May 1902 (M. A.
Carriker), 2cJ, 19; 22 Aug. 1927 (Lankester and Rehn), 3<J,
19. Between La Union and Buenos Aires, Terraba Valley,
5500 ft. [1670 m], May 1935 (Lankaster), 1 9 . Juan Vinas, 3300
ft. [1000 m], Mar. 1902 (L. Beamer), 1 6* , 1 9 ; 27 Jun. 1909 (P.
P. Calvert), IcJ, 19. Peralta, 8 Aug. 1909 (Calvert), IcJ; May
1923 (Lankester), Id. La Emelia near Guapiles, Aug. 1923,
Sep. 1927 (Rehn), 4d. Cariblanca, 600 m (Lankester), 19.
Parisimina, 5 m, 26 Jul. 1928 (M. Valerio), 19. PUNTARENAS:
Rio Cataratas, near Brujo, Sep. 1979 (Rowell), 16, 19. ALA-
JUELA: 5 km s San Lorenzo, Sep. 1979 (Rowell), Id, 19,
CHFR.

Nicaragua. CHONTALES: (Janson), 13, 19 (poorly pre-
served but appears to be this species).

COMMENTS. — Although we have no records
of C. trogon trogon and C. psittacus occurring
together, it seems possible that they do. For ex-
ample, we have this species from near Guapiles
and C. psittacus from Siquirres about 30 km dis-
tant in similar lowland forest country.

7. Chromacris trogon intermedia n.subsp.

DIAGNOSIS. — Wing orange with no black on
basal half of first two plaits, or first paired fold
as in C. color ata. Antennal segments all black.
Hind femur with yellow bands that may be
weakly or strongly developed. Hind tibia with
no bands. Prominent wide yellow stripe along
ventral margin of lateral lobe of pronotum that

extends onto cheek of head. Prosternal tubercle
short, tapering rapidly to a point. Distinguished
from C. psittacus by lack of banding on hind
tibia, and from the nominate subspecies of C.
trogon by banding on hind femur (Table 2).

DISTRIBUTION. — Northern Guatemala, Be-
lize, and Honduras.

SPECIMENS. — Holotype: 6, ANSP; Honduras, Lancertilla
nearTela, Dept. Atlantica, 100-800 ft. (30-250 m), rainforest,
8 Nov. 1930.

Paratypes: Same data as for holotype, 3 9 . Belize. 50 mi. [80
km] s El Cajo, Mountain Pine Ridge road, 17 Aug. 1960 (P.
N. Litchfield), Id, 29, UMMZ. Guatemala. Piedras Negras,
600-800 ft. [180-240 m], 30 Jun. 1933 (D. W. Amran), 1 9 .

COMMENTS. — This subspecies is intermediate
between C. trogon trogon and C. psittacus in
the reduction in banding of the hind leg, which
might suggest that they should be treated as
three subspecies, but C. trogon trogon and C.
psittacus occur close together in Costa Rica,
though as noted under the subspecies, C. trogon
trogon, they have not been recorded from the
same locality. Specimens from Nicaragua and
Honduras are needed to help clarify the prob-
lem.

8. Chromacris psittacus psittacus (Gerstaecker)

Romalea psittacus GERSTAECKER, 1873:185. [Lectotype, d,
among four male syntypes with same data, one marked
"typus" and here designated; Berlin Museum; Bogota, Co-
lombia. Species of Chromacris probably do not occur in the
vicinity of Bogota. ANSP has specimens of C. icterus also
labelled Bogota, and this species actually occurs at lower
elevations on the eastern slopes of the Andes.]

ROBERTS AND CARBONELL: REVISION OF CHROMACRIS AND XESTOTRACHELUS

FIGURES 26 and 27. Internal male genitalia of Chromacris miles from Quintana Roo, Mexico (Fig. 26), and Xestotrachelus
robustus from Ihu, Caaguazvi, Paraguay (Fig. 27). A, phallus, lateral; B, cingulum, lateral; C, endophallus, lateral; D, phallus,
dorsal; E, cingulum, dorsal; F, endophallus, dorsal; G, epiphallus, dorsal; H, epiphallus, frontal; /, epiphallus, lateral, left side.

DIAGNOSIS. — Wings yellow to orange-yellow,
lacking black on anterior basal half (Fig. 8). An-
tennae all black. No yellow on dorsum of me-
tazona of pronotum. Hind femur with three yel-
low bands, but proximal band not entire. Hind
tibia with two bands. For comparison with other
species see Table 2.

SPECIMENS. — Costa Rica. Siquirres, 3 Jul. 1903 (M. A. Car-
riker), 7d, 19. Ujarass de Terraba, 10 Sep. 1907, Id. Monte
Verde, "summer," 1928 (F. G. Wallace), Id, 12. Castilla
Farm, lower Rio Reventazon, 29 Jul. 1936 (C. W. Dodge), Id .

Panama. Gatiin, Jul.-Aug. 1916 (D. E. Harower), 8d, 32.
Cabima, 24 May 1911 (August Busck), 2<J, 1 juv. Barro Col-
orado Island, C.Z., 22 Jul. 1933 (H. H. Hood), 1 2 .

Colombia. CUNDINAMARCA: Las Mesitas, Sep. 1915 (A.
Maria), Id, 32.

Venezuela. ZULIA: Kasmera, Rio Yasa, Sierra de Perija, 250
m, 19 Sep. 1961, 26, 32, Universidad Central, Institute de
Zoologia Agricola, Maracay.

COMMENTS. — Evidently seasonal. One of us
(H.R.R.) visiting Costa Rica for several years in
the Pacific and Caribbean lowlands during Feb-
ruary and March never encountered this species
or C. trogon. Most of our records suggest that
it occurs commonly May to September.

9. Chromacris psittacus pacificus n.subsp.

DIAGNOSIS. — Similar to C. psittacus psittacus
except proximal yellow band on lower portion

of hind femur entire rather than interrupted. See
Figure 1 and Table 2.

SPECIMENS. — Holotype: d , ANSP; Ecuador, Dos Puentes,
below Naranjapata along the Guayaquil-Quito railway in
Chanchan River valley, 1750 ft. [530 m], 15 Mar. 1931 (W. J.
Coxey). [Additional information on location of Dos Puentes
is added here from Coxey 1927:10.]

Paratypes: Colombia. EL VALLE: Jimenez, 1600 ft. [486 m],
19 Mar. 1907 (M. G. Palmer), 2d , 1 9 ; Choco (M. G. Palmer),
1$. ANTIOQUIA: Andagoya, 12; Cordillere, "vers occid. Rio
Yurumaqui," 1933 (E. Aubert de la Rue), Id, 12, MNHN.
NARINO; "entre Guayacana et el Diviso," 80 m, Nov. 1968
(M. Descamps), Id.

Ecuador. CHIMBORAZO: Dos Puentes, 1750 ft. [533 m], 11
Jan. 1921 and 15 Mar. 1931 (W. J. Coxey), 5d, 22; Ventura,
1400 ft. [469 m], 10-13 Apr. 1922, 3d, 32. GUAYAS: Bucay,
900 ft. [274 m], 19 Mar. 1922 (G. H. Tate), 1 2 . TUNGURAHUA:
Ambato, Id, MNHN; Balzapamba, near Ambato (R.
Haensch S.), Id, Berlin Museum.

COMMENTS. — The slight but consistent differ-
ence in the form of the proximal yellow band on
the hind femur of these specimens warrants sub-
specific recognition.

10. Chromacris icterus (Pictet and Saussure)

Rhomalea icterus PICTET AND SAUSSURE, 1887:353. [Lecto-
type, 2 , so labelled and here designated, Geneva Museum;
Quito, Ecuador.]

Rhomalea opulenta GERSTAECKER, 1889:32. [Holotype, 2,
Zoological Museum, University of Greifswald; Sao Paulo

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4

de Olivenca, Amazonas, Brazil. Holotype examined. New
synonym.]

DIAGNOSIS. — Tip of antenna yellow. Anterior,
basal portion of hind wing entirely yellow (Fig.
9). Yellow patches on dorsum of metazona of
pronotum. Three yellow bands on hind femur.
Single proximal band on hind tibia distinguishes
species from all others (see Table 2). A large
species with relatively long wings; males 30-38
mm, females 40-55 mm from fastigium to wing
tips.

DISTRIBUTION. — Southeastern Colombia,
eastern Ecuador, northwestern Brazil and
northeastern Peru.

SPECIMENS. — Colombia. BOGOTA: (so labelled but probably
in error) (A. Maria), 26, 49. CUNDINAMARCA: Susumuco,
Feb., Sep. 1916, 1917 (A. Maria), 3d, 29. META: Villavicen-
cio, May, Jun. 1919 (A. Maria), 26 , 29. PUTUMAYO: "bord
riviere Mocoa," 800 m, Nov. 1968 (M. Descamps), Id,
MNHN.

Ecuador. MORONA-SANTIAGO PROV.: s of Mendez, 800 m,
19-21 Oct. 1977 (L. E. Pena), 19.

Peru. LORETO: Putumayo District, La Chorrera to La Som-
bra, 21 Aug. 1920, 1 9 ; Iquitos, 8 Jan. 1920 (H. S. Parish), 1 9 .

Brazil. AMAZONAS: "Hyntanahan" probably Huitanaa, Rio
Purus, Jan. 1922 (S. M. Klages), 1 9 ; Tabatinga, Sep.-Dec.
1977 (L. C. Pereira, B. Silva), long series both sexes, CACS;
Eirunepe, Jun. 1950 (J. C. Carvalho), 26 , 1 9 , CACS; Atalaya
do Norte, Nov. 1977 (B. Silva), long series, CACS.

1 1 . Chromacris peruviana (Pictet and Saussure)

Rhomalea peruviana PICTET AND SAUSSURE, 1887:352. [Lec-
totype, 9 , so labelled by C.S.C. and here designated from
among Id and 39 syntypes, Geneva Museum; Peru.]

DIAGNOSIS. — Similar to C. icterus in having
yellow wings and yellow antennal tips, but dif-
fers by having two yellow bands on hind tibiae
rather than one (Table 2); differs from all other
yellow-winged species by having the black on
anterior portion of first plait of hind wings ex-
tend to base of wing (Fig. 10). Tegmina relative-
ly long, narrow, and greatly surpassing ends of
hind femora; anterior and posterior margins bor-
dered by yellowish or pale green areas. This teg-
minal coloration is unique for genus.

SPECIMENS. — Peru. JUNIN: Chanchamayo, a district around
La Merced in valley below Tarma, 2000-3000 ft. (610-914 m),
1 d ; Satipo, near Huancayo, 1650 m, Jul. 1844 (Schunke), 1 d ;
Mar., Jun. 1944 (P. Paprzycki), 3 9 ; Puerto Yessup, Feb. 1930
(M. A. Carriker), 19. HUANUCO: Leonpampa, 110 km E
Huanuco, Dec. 1937 (Felix Woytkowski), 3d , 1 9 ; Tingo Mar-
ia, 670 m, Sep. 1946 (Weyrauch), 39; same locality, 2 Dec.
1954 (E. D. Schlinger, E. S. Ross), 19; Divisoria, Cordillera
Azul, 1500 m, Id, 19. LORETO: Rio Aguaytia, between Tingo
Maria and Pucallpa, 400 m, Feb. 1961, Id , 29 . SAN MARTIN:
Prov. Huallaga, Rio Mixiollo, 1200 m, 7 Aug. 1900 (C. A.

Baer), 19. Cuzco: Paucartambo, Pilcopata, Nov. 1968 (F.
Carrasco), Id, 19; Paucartambo, Atalaya, May 1976 (Des-
camps, Carbonell), Id, 19, CACS.
Venezuela. ARAGUA: Nov. 1942, 39.

COMMENTS. — The Venezuelan specimens are
undoubtedly this species. However, C. icterus
occurs between this and the Peruvian localities,
suggesting the need to confirm the correctness
of the locality.

Xestotrachelus Bruner

Xestotrachelus BRUNER, 1913:469. [Type-species Xestotrach-
elus hasemani Bruner (=X. robustus) by original designa-
tion.]

DIAGNOSIS. — Red and black pattern of hind
wings (Fig. 11) closely resembles red-wing
species of Chromacris, but head and thorax are
much more robust, and tegmina may extend well
short of, or only slightly beyond, end of hind
femora (Fig. 2). Head, pronotum, and other
parts of body and appendages may be contrast-
ingly marked with black, pale olive-yellow, and
red. Hind tibiae usually red. Phallic structures
described under X. robustus.

DISTRIBUTION. — Known from Maranhao in
northeastern Brazil, Bahia, Espfrito Santo,
Mato Grosso; Paraguay; and eastern Bolivia.

12. Xestotrachelus robustus (Bruner)

Zoniopoda robusta BRUNER, 1911:60. [Lectotype, d, here
designated, ANSP; Chapada dos Guimaraes, Mato Grosso,
Brazil. The type-series consisted of a male and female, each
labelled as the type.]

Xestotrachelus hasemani BRUNER, 1913:470. [Holotype, 9,
ANSP; labelled as from Galhao, not Calhao as reported by
Bruner, Rio Sapao, western Bahia, Brazil. Actually Galhao,
10°35'S, 46°15'W, is in Goias on the Rio Galhao, and the
Rio Sapao is nearby in Bahia. New synonym.]

DIAGNOSIS. — See diagnosis of genus. Extent
of black on prozona and mesozona of pronotum
variable, may be entirely black, or divided to
form two black transverse bands. Extent of red
on sides and undersides of abdomen also van-,
able. Easily distinguished from species of Chro-
macris by red hind tibia and lack of banding on
hind femur.

Phallic structures generally similar to those in
Chromacris. Aedeagal valves (Fig. 21) strongly
sclerotized, much larger, and sculptured. Rami
of cingulum (Fig. 275) narrow, partly surround-
ing base of aedeagus, strongly bent inward and
ventrad near their end, and outer surface at bend
covered with small spines. Epiphallus (Fig.

ROBERTS AND CARBONELL: REVISION OF CHROMACR1S AND XESTOTRACHELUS

27G-7) similar to that in Chromacris except that
the ancorae are well developed.
DISTRIBUTION. — See distribution of genus.

SPECIMENS. — Brazil. MARANHAO: Barra do Corda, Feb.
1955, 19, CACS. BAHIA: Maracas, Feb. 1963 (F. M. Oliv-
eria), 39, CACS. ESPI'RITO SANTO: Linhares, Mar. 1981 (B.
Silva), Id, CACS. GOIAS: in addition to the type-locality of
X. hasemani, 60 km w Mineiros, 10 Mar. 1980 (Roppa, Car-
bonell, Roberts), Id. MATO GROSSO: in addition to the type
of X. robustus, Corumba, Urucum, 23-29 Dec. 1919 (R. G.
Harris), 4 juv., 18d, 149. MATO GROSSO SUL: 30-60 km E
Aquidauana, 16 Mar. 1980 (Roppa, Carbonell, Roberts), Id.

Bolivia. SANTA CRUZ: prov. of Sara, 450 m, Jan.-Feb. 1922
(J. Steinbach), 8d , 169 ; Buena Vista, 500 m, 3 Feb. 1922 (J.
Steinbach), 19.

Paraguay. CAAGUAZU: near Ihu, Mar. 1965 (Carbonell,
Mesa, Monne), Id, 29, CACS. AMAMBAY: Cerro Cora, Jan.
1972 (Descamps, Ronderos, Carbonell), 8d, 59, 1 last instar
nymph, CSC.

COMMENTS. — Individual geographic variation
in relative tegminal length is evident. In the
series recorded from eastern Bolivia, tegmen
shorter than or about as long as hind femur. In
the series from Corumba, tegmen slightly or de-
cidedly longer than femur. Tegmen of male from
near Aquidauana 19 mm and hind femur 15.5
mm in length, whereas in the male from Minei-
ros, Goias, tegmen 16 mm and femur 17 mm.
Tegmen of female holotype of X. hasemani is
26 mm, hind femur 18 mm. This specimen has
a decidedly longer tegmen in proportion to the
hind femur than do specimens to the south. It
seems possible, however, that populations to be
found between this type-locality and those pres-
ently observed to the south will be intermediate
in relative tegminal length. The aedeagus of the
male from Espirito Santo is relatively shorter
and therefore appears to be somewhat broader
in lateral view than others examined. The red
patch near the end of the hind wing in the longer-
winged specimens from Linhares and Maranhao
connects narrowly with the large red field of the
wing. Conceivably, this modification is the re-
sult of the lengthened wing. With more evidence
the longer-winged hasemani form might be rec-
ognized as a subspecies.

Noteworthy is the last instar nymph from Cer-
ro Cora, Paraguay, recorded above, the color of
which is well preserved and matches the color
pattern of the adult. Unlike the species of Chro-
macris which have a bright but relatively cryptic
coloration in the adult stage, X. robustus has a
much more striking coloration in the adult,
which may well serve as a warning to predators.

Assuming this is correct, then there would be no
need to develop a different coloration in the
nymphal stages as in Chromacris.

LITERATURE CITED

AMEDEGNATO, C. 1974. Les genres d'acridiens neotropicaux,

leur classification par families, sous-families et tribus. Ac-

rida 3: 193-204.
ASTACIO-CABRERA, O. 1975. Notas sobre algunas acridioi-

deos de Nicaragua. Organismo Internacional Regional San-

idad Agropecuaria, Managua, Nicaragua. 41 p.
BARRERA, M., AND S. Z. TURK. 1977. Acridios del NOA. II.

Contribution al conocimiento de huevos, desoves y habitos

de postura de algunas especies de tucuras (Orthoptera, Ac-

rididae) de la Provincia de Tucuman. Acta Zoological Lil-

loana 32(9): 167-188.
BRUNER, L. 191 1. South American Acridoidea. Annals of the

Carnegie Museum 8(1):5-I47.
. 1913. South American locusts (Acridoidea) II. Annals

of the Carnegie Museum 8(3-4): 423-506.
CARRASCO-Z, F. 1962. Observaciones sobre algunas plagas

de interes para la zona del Cuzco. Revista Peruana Ento-

mologia Agricola 5:97-100.
COXEY, W. J. 1927. Impressions of Ecuador. Year Book

(1926) Academy of Natural Sciences of Philadelphia: 5-20.
DRURY, D. 1773. Illustrations of natural history, wherein are

exhibited upwards of two hundred and twenty figures of

exotic insects, etc., vol. II. London. 92 p., 50 pi.
GAUGLIUMI, P. 1973. Pragas de cana-de-acucar, nordeste do

Brazil. M.I.C. Institute do A9ucar e do Alcool. Cole9§o

Canavieira, no. 10. Rio de Janeiro, v + 622 p.
GERSTAECKER, A. 1873. Acridiodea nonulla nova insigniora.

Entomologische Zeitung, Stettin 34(1-3): 185-197.
. 1889. Charakteristik einer Reihe bemerkenswerther

Orthopteren. Mittheilungen aus dem naturwissenschaftlich-

en Verein Neu-Pommern und Riigen in (Greifswald) Berlin

20:1-58.
HAHN, C. W. 1835. Icones Orthoptorum. Nurnberg. 3 p., 4

pi.
KIRBY, W. F. 1910. A synonymic catalogue, Orthoptera Sal-

tatoria. British Museum, London 3(2): 1-674.
PICTET, A., AND H. DE SAUSSURE. 1887. Catalogue

d'Acridiens. Bulletin de la Societe Entomologique Suisse

7(9):33 1-376.
PRETTO-MALCA, R. 1968. Estudios del ciclo biologica, mor-

fometrfa y etiologi'a de Chromacris colorata (Serville), (Or-
thoptera, Acridoidea). Institute Tecnologia, Monterrey,

mimeographed report: 83-87.
RAGGE, D. R. 1955. The wing-venation of the Orthoptera Sal-

tatoria. British Museum (Natural History) London. 159 p.
REHN, J. A. G. 1904. Notes on Orthoptera from northern and

central Mexico. Proceedings of the Academy of Natural

Sciences of Philadelphia 31:513-548.
, AND H. J. GRANT. 1959. An analysis of the tribes of

the Romaleinae with special reference to their internal gen-

italia (Orthoptera: Acrididae). Transactions of the American

Entomological Society 85:233-271.
ROWELL, H. F. 1978. Food plant specificity in neotropical

rain-forest acridids. Entomologie Experimentia et Applicata

24:451-462.
SAUSSURE, H. DE. 1859. Orthoptera nova Americana. Revue

Magazin Zoologie (2)1 1:390-394.

58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 4

SERVILLE, M. A. 1839. Histoire Naturelle des Insects Or- Estudios biologicos, morfometricos y aspectos ecologicos

thopteres, Paris: 1-776. de Chromacris speciosa (Thunberg) (Acrididae, Romalei-

THUNBERG, C. P. 1824. Grylli Monographia, illustrata. — nae). Acta Zoologica Lilloana 32(6): 12 1-146.

Memoirs de 1' Academic Imperiale des Sciences de St. Pe- WALKER, F. 1870. Catalogue of the specimens of Dermaptera

tersbourg 9:390-430. Saltatoria in the collection of the British Museum 4:605-

TURK, S. Z., AND M. BARRERA. 1976. Acridios del NOA. I. 809.

CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, California 941 18

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 5, pp. 59-66, 5 figs., 1 table.

November 4, 1982

A NEW GENUS AND TWO NEW SPECIES OF

REMARKABLE PACIFIC WORM EELS
(OPHICHTHIDAE, SUBFAMILY MYROPHINAE)

John E. McCosker

Steinhart Aquarium, California Academy of Sciences, Golden Gate Park,
San Francisco, California 94118

ABSTRACT: Glenoglossa wassi gen. et sp.nov., described from Samoa, differs from all other myrophine
ophichthids in its elongate tongue decorated with a lure, and in certain osteological and cephalic pore con-
ditions. Neenchelys daedalus sp.nov., is described from midwater-captured juveniles and adults from off New
Guinea and the central Pacific and represents the second known midwater worm eel; it differs from its
congeners in its extreme elongation and vertebral number. The status of the species of Pseudomyrophis and
Neenchelys is discussed. An analytical key to the genera of the ophichthid subfamily Myrophinae is provided.

INTRODUCTION

The snake eels and worm eels of the family
Ophichthidae are the most diverse and inventive
of true eels. The more than 220 species distrib-
uted among more than 53 genera inhabit all trop-
ical and subtropical oceans and seas, and have
invaded the intertidal zone, coral reefs, shallow
substrates, and even the midwater realm. To this
array of astounding evolutionary forays, I add
two remarkable new western Pacific species, one
representing a distinct new genus of worm eels,
of the subfamily Myrophinae (sensu McCosker
1977). The first, Glenoglossa wassi gen. et sp.nov.,
is unique among eels in having modified its glos-
sohyal into a lure, not unlike that of a urano-
scopid stargazer or an antennariid. The other,
Neenchelys daedalus sp.nov., represents a sec-
ond, independent invasion of the midwater realm
by an ophichthid.

MATERIALS AND METHODS

Measurements are straight-line, made either
with a 300 mm ruler with 0.5 mm gradations
(for total length [TL], trunk length, and tail length)
recorded to the nearest 0.5 mm, or with dial
calipers (all other measurements) and recorded
to the nearest 0.1 mm. Body length comprises
head and trunk lengths. Head length (HL) is mea-
sured from the snout tip to the posterodorsal
margin of the gill opening; trunk length is taken
from the end of the head to mid-anus; maximum
body depth does not include the median fins.
Vertebral counts, which include the hypural, were
taken from radiographs. Stained and cleared
specimens were prepared using the Taylor (1967)
trypsin technique. Institutional abbreviations of
material examined are explained in the
Acknowledgments.

[59]

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 5

ANALYTICAL KEY TO THE GENERA OF
OPHICHTHIDAE, SUBFAMILY MYROPHINAE

la. All branchiostegal rays originate either in
association with hyoid or before level of
epihyal tips; free rays, when present, fewer
than attached; tail tip a hard or fleshy fin-
less point; gill opening midlateral to

entirely ventral, unconstricted

subfamily Ophichthinae

Ib. Accessory branchiostegal rays originate
behind ends of epihyal, free rays more
numerous than attached; caudal fin rays
conspicuous, confluent with dorsal and
anal, tail tip flexible; gill openings mid-
lateral, a constricted opening (subfamily
Myrophinae) 2

2a. Anterior nostril non-tubular, posterior
nostril before eye; eye large, ca. 6 times
in head length; pectoral fin moderately
developed Benthenchelys Fowler, 1934

2b. Anterior nostril tubular, posterior nostril
either before eye, along upper lip, or within
mouth; eye smaller, 10 or more in head;
pectoral fin may be absent 3

3a. Posterior nostril before eye, above the lip
and not covered by a flap; pectoral fin
present, but may be reduced to a small,
barely noticeable flap in posterodorsal
corner of gill opening 4

3b. Posterior nostril labial, either within lip
and opening into mouth, or along lip and
covered by a flap; pectoral fin either pres-
ent and well developed or absent 5

4a. Dorsal fin origin in anterior trunk region;
snout conical; pectoral fin well developed,
=s snout; third preoperculomandibular

pore (pop3) absent

Neenchelys Bamber, 1915

4b. Dorsal fin origin in posterior trunk region;
snout broad, tumid; pectoral fin minute,

< eye; pop3 present

Pseudomyrophis Wade, 1 946

5a. Pectoral fin well developed; pleural ribs
absent behind 1 5th-20th vertebrae 6

5b. Pectoral fin absent; pleural ribs present on
all trunk vertebrae 7

6a. Dorsal fin origin above or behind anus;
maxilla stout, not tapering posteriorly, and
abutting pterygoid; vomerine teeth absent
Ahlia Jordan and Davis, 1891

6b. Dorsal fin origin anterior to mid-trunk
region; maxilla thin and tapering poste-
riorly, not closely associated with ptery-
goid; vomerine teeth present

Myrophis Liitken, 1851

7a. Tongue elongate, extending well beyond
mouth and decorated with a fleshy appen-
dage; inner edge of lips and palate deco-
rated with fleshy lappets; teeth conical and
uniserial Glenoglossa novum

7b. Tongue not elongate, not extending out-
side of mouth, lacking a fleshy appendage
at its tip; inner edge of lips and palate
smooth; teeth either conical or blunt, uni-
serial or multiserial 8

8a. A prominent median toothed groove on
ventral side of snout, bordered by dermal
folds, extends anteriorly to anterior nos-
trils; anterior nostrils elongated tubes equal

to eye in length

Schismorhynchus McCosker, 1970

8b. Ventral side of snout without a prominent
median groove bordered by dermal folds;
anterior nostrils less than eye in length .... 9

9a. Teeth absent on vomer, absent or embed-
ded on intermaxillary, those on maxillary
and dentary minute or villiform; dorsal

fin origin behind anus

Schultzidia Gosline, 195 1

9b. Teeth present on intermaxillary, maxil-
lary, dentary, and vomer; dorsal fin origin

either before or behind anus

Muraenichthys Bleeker, 1853

Glenoglossa McCosker, gen.nov.

TYPE-SPECIES. — Glenoglossa wassi McCosker, sp.nov.

DIAGNOSIS.— Body moderately elongate, lat-
erally compressed posteriorly; snout conical,
grooved on underside; anterior nostril within a
tube, posterior nostril within a short tube at outer
edge of lip, directed ventrally; dorsal fin origin
slightly before anus; pectoral fin absent; pop3
absent; tongue elongate, extends beyond mouth,
decorated with fleshy appendage; inner edge of
lips and palate decorated with fleshy lappets; teeth
conical, uniserial, absent on vomer; gill arches
reduced, third hypobranchial and second infra-
pharyngobranchial absent, third and fourth upper
pharyngobranchial tooth plates weakly fused;
suspensorium nearly vertical, pterygoid slender

McCOSKER: NEW WORM EELS

FIGURE 1. Holotype of Glenoglossa wassi McCosker, sp.nov., CAS 47049, 153.5 mm TL. Arrows indicate origin of dorsal
and anal fins.

and reduced; cleithrum and supracleithrum
reduced to thin slivers. Other characters those of
the single species.

ETYMOLOGY.— From the Greek yXf/voa (gle-
nos), a thing to stare at, and y\&aaa (glossa, fem-
inine), tongue.

Glenoglossa wassi McCosker, sp.nov.

(Figures 1-3)

HOLOTYPE.— CAS 47049, 153.5 mm TL, collected using
rotenone over sand at base of large coral head, 40 m depth,
Larsen Bay, Tutuila Island, American Samoa; R. Wass and G.
Yamasaki, 18 Nov. 1975.

PARATYPE.— CAS 47048, 88.8 mm TL, collected using rote-
none along sloping sand bottom at base of coral and lava cliff,
40 m depth, Steps Point, Tutuila Island, American Samoa; R.
Wass and R. Lubbock, on 4 Mar. 1975. This specimen was
cleared and stained.

COUNTS AND MEASUREMENTS.— The condition
of the holotype is parenthetically followed by
that of the paratype: TL 1 53.5(88.8); head length
20.8(12.8); trunk length 54.7(30.0); tail length
78.0(46.0); body depth at gill openings 5.6(3.7);
body width at gill openings 3.0(1.9); origin of
dorsal fin 67. 1(39.6); snout length 3.5(2.3); upper
jaw length 6.7(4.4); eye diameter 1.1(0.7); inter-
orbital distance 2.0(1.0). Total vertebrae
127(126), preanal vertebrae 54(53).

DESCRIPTION. — Body moderately elongate,
depth at gill oepnings 24-26.5 in TL, tapering
and laterally compressed posteriorly. Head and
trunk 2.03-2.07 and head 6.9-7.4 in TL. Snout
acute; lower jaw included, tip slightly in advance
of anterior nostril base. Anterior nostril tubular;
posterior nostril at edge of lip, entirely outside
of mouth, within small tube. Eye slightly in

advance of midpoint of upper jaw. Gill opening
mid-lateral, a constricted opening.

Median fins low, lying partially within a groove,
meeting each other and extending noticeably
beyond caudal tip. Dorsal fin arises less than a
head length in advance of anus.

Head pores minute, difficult to discern. Single
temporal and interorbital pores. Four pores along
mandible. Two preoperculomandibular pores.
Lateral line pores difficult to identify in pre-
served specimens; approximately 1 1 pores before
the gill opening.

Tongue extends from mouth. A fleshy appen-
dage, differing slightly in each specimen (Figs. 2-
3), extends beyond the slender glossohyal. The
inner edge of lips, floor of mouth, and palate
flanked by fleshy tissue (Fig. 2).

Teeth small, conical, uniserial in jaws. An
intermaxillary chevron of four teeth, followed by
two medial teeth. Vomerine teeth absent. Nine
teeth along maxilla, 1 4 along mandible.

Body color in isopropyl alcohol uniformly tan.
Numerous, minute brown punctations in mouth,
along head and dorsal body surface. Fins pale.
Base of lure has a dark spot. Eyes dark blue. Color
of paratype in life, recorded by R. Wass, "light
greenish-yellow with tiny purple brown specks.
Lure transparent with black 'eye.' "

ETYMOLOGY.— Named in honor of Richard C.
Wass, collector of these and many other impor-
tant fishes from Samoa.

REMARKS.— This myrophine is remarkable in
the development of its tongue which, because of
its length and appearance, serves as a lure to
attract small fish. It is the only eel known to use

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 5

FIGURE 2. Open mouth of holotype of Glenoglossa wassi
McCosker, sp.nov., CAS 47049.

such a feeding strategy, and is certainly no less
remarkable than those of certain uranoscopids,
ceratioids, or the alligator snapping turtle. The
"eyes" and appendages of the lure would indicate
that it resembles a small crustacean. In the stom-
ach of the holotype was a partially digested goby-
like fish nearly 2 cm in length. The other oral
decoration provided by the fleshy labial lappets
suggests that the eel lies buried in the sand with

FIGURE 3. Tongue lure of paratype of Glenoglossa wassi
McCosker, sp.nov., CAS 47048.

its head exposed, luring prey by flicking its glos-
sohyal.

Glenoglossa wassi is most closely related to
species of Muraenichthys and Schismorhynchus.
It is easily separable from them on the basis of
its tongue development, its absence of vomerine
teeth, and certain other osteological characters.
It is most like Schismorhynchus in its general
facies, the development of its snout groove,
reduced pterygoid, and gill arch reductions, con-
ditions which probably relate to the feeding
behavior of the species involved. It is clearly a
specialized worm eel whose ancestry is in the
sharp-snouted, conical-toothed species group of
Muraenichthys (sensu McCosker 1977) that gave
rise to Schismorhynchus and may have shared a
common ancestor with it.

Neenchelys Bamber, 1915

TYPE-SPECIES.— Neenchelys microtretus Bamber, 1915.

DIAGNOSIS. — Body moderately to extremely
elongate, laterally compressed behind head; tail
much longer than head and trunk; snout conical,
anterior nostril in a tube, posterior nostril an
elongate slit entirely before eye; eye moderate;
dorsal fin origin mid-trunk; pectoral fin moder-
ately developed, girdle limited to reduced clei-
thrum and supracleithrum; gill opening reduced,
a constricted hole; pop3 absent; teeth few, conical
and slender, uniserial on jaws and vomer; max-
illary attachment at mid-vomer; gill arches
reduced; suspensorium nearly vertical, slightly

McCOSKER: NEW WORM EELS

FIGURE 4. Holotype of Neenchelys daedalus McCosker, sp.nov., AMS 1.19690-012, 341.5 mm TL.

inclined anteriorly; pterygoid slender and
reduced; neurocranium rounded, lacking a crest;
color uniform.

Neenchelys daedalus McCosker, sp.nov.

(Figures 4-5)

HOLOTYPE. -AMS 1.19690-012 (field no. JP 69-29), held in
trust for Papua New Guinea National Fish Collection, 341.5
mm TL, a female with eggs, captured in Astrolobe Bay, s of
Madang, Papua New Guinea (5°24'S, 145°52.5'E), 6-ft Isaacs-
Kidd midwater trawl (IKMT), 0-140 fms [0-256m] over a
"rough peak 500+ fms" bottom, 1850-2100 h, 7 Oct. 1969.
In that same collection were numerous myctophids, gono-
stomatids, and chauliodontids.

PARATYPE.-CAS 50708, 272.9 mm TL, collected with the
holotype.

COUNTS AND MEASUREMENTS (in mm).— The
condition of the holotype is parenthetically fol-
lowed by that of the paratype: TL 341.5(272.9);
head length 23.5(2 1 .4); trunk length 65(53.5); tail
length 253(198); body depth at gill openings
6.6(5.9); body width at gill openings 4.7(4.0); body
depth at anus 6.2(5.0); body width at anus
4.2(3.5); origin of dorsal fin 49.5(42); length of
pectoral fin 5.2(5.0); snout length 3.9(3.8); upper
jaw length 6.2(5.2); eye diameter 1.2(1.2); inter-
orbital distance 2.05(1.8). Total vertebrae
235(225); preanal vertebrae 59(58).

DESCRIPTION.— Body extremely elongate, depth
at gill opening 46.3-5 1 .7 in TL, tapering slightly
and laterally compressed posteriorly. Tail much
longer than head and trunk, 1.34-1.38 in TL.
Head and trunk 3.64-3.86 and head 12.8-14.7
in TL. Snout moderately acute; lower jaw
included, its tip extends to anterior base of ante-
rior nostril. Top and sides of snout and edge of
lower jaw covered with numerous small papillae

(presumably sensory in function), slightly smaller
than cephalic pores. Anterior nostril tubular,
directed ca. 45° laterally, posterior nostril an
elongate slit above upper lip, nearly as wide as
orbit. Eye begins behind midpoint of jaw, mod-
erate in size, 4.3-5.2 in jaw length. Gill openings
low on side, each a constricted opening.

Pectoral fin well developed, nearly as long as
gape. Median fins well developed and obvious,
not lying in a groove as in most myrophines.
Anal fin larger than dorsal, nearly as deep as
body. Dorsal fin arises about mid-trunk, the pre-
dorsal distance 6.49-6.89 in TL.

Head pores small but apparent (Fig. 5). Single
temporal and interorbital pores. Five mandib-
ular pores, and two over preopercle. Two post-
orbital pores. Lateral line pores small but obvious;
14 on head, 61 before anal opening, not discern-
ible in posterior tail region.

Teeth few, slender, conical, of moderate size
for a myrophine. A single premaxillary tooth,
flanked by a pair of retrorse teeth, followed by
two medial teeth, then a pair of teeth, at which
point the maxillae attach, each possessing 6-7
uniserial teeth. Vomer has three teeth that end
about midway along toothed portion of maxilla.
Lower jaw teeth uniserial, 17-18 on each side.

Gill arches reduced; first basibranchial absent,
third and fourth infrapharyngobranchial tooth
plates weakly fused. Branchiostegal rays numer-
ous, unbranched; eight attached to hyoid (1 along
the ceratohyal, 7 along the epihyal), 25 unat-
tached, on each side.

Body color in isopropyl alcohol uniform tan,
except belly which is dark brown to black. Fins
colorless. Eye dark blue.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 5

FIGURE 5. Head of holotype of Neenchelys daedalus McCosker, sp. nov., AMS 1. 1 9690-0 1 2.

ETYMOLOGY.— Named daedalus, a noun in
apposition, in honor of the Greek artisan who
escaped from his Earth-bound prison and
ascended into heaven.

REMARKS.— All specimens of Neenchelys dae-
dalus have come from midwater. It therefore
seems likely that it is a midwater eel, although
the possibility exists that juvenile and subadult
specimens were merely transformed leptocephali
that were captured just prior to settlement, and/
or the adults were benthic eels that were captured
en route to a surface spawning event. Both pos-
sibilities are unlikely in that the size range of the
specimens is too great and their condition is too
"uniform" to have been captured during trans-
formation, and none appear to have retained lar-
val conditions. Furthermore, the morphometric
changes in eye size undergone by surface-mi-
grating benthic myrophines, such as Ahlia
egmontis (see Cohen and Dean 1970, and
McCosker 1977), are absent. Further evidence
for a midwater habitat is provided by the par-
tially digested crustacean material in the gut of
the cleared-and-stained specimen (which sug-
gests that it had fed prior to capture and not while
in the net) as well as the darkened vent and pres-
ence of sensory papillae on the snout.

The evidence thus suggests that N. daedalus is
the second ophichthid known to have left the

substrate to adopt a midwater life style. This
adaptation has been independently achieved,
however, in that its closest relatives are benthic,
fossorial species. The other midwater ophichthid,
Benthenchelys cartieri Fowler, lives pelagically
at 100-250 m over deep water in the central
Indo-Pacific (Castle 1972) and displays many
similar adaptations, such as enlarged median fins,
sensory papillae, an elongate tail, and slender
teeth.

The new species is more similar in appearance,
owing to its extreme elongation, to the Atlantic
species of Pseudomyrophis than to its congeners.
However, its osteology, pore condition, and fin
size and placement are in agreement with
Neenchelys microtretus Bamber, 1915, N. bui-
tendijki Weber and de Beaufort, 1916, and the
sketchily described N. parvipectoralis Chu, Wu,
and Jin, 1981. Castle (1980) has illustrated the
larvae of Neenchelys and commented upon their
distribution. The new species may be separated
from its congeners using the characters in Ta-
ble 1.

In an earlier publication (McCosker 1977), I
cautiously recognized the generic distinction
between Neenchelys and Pseudomyrophis. My
subsequent examination of additional osteolog-
ical preparations of Neenchelys spp. and Pseu-
domyrophis spp. have further substantiated those

TABLE 1. VERTEBRAE AND BODY PROPORTIONS (in thousandths of TL) OF THE SPECIES OF NEENCHELYS.

Head

Tail

Body

Total

length

depth

vertebrae

N. microtretus

114'

580

38'

1512

N. buitendijki

117-127'

565-643

31-51'

142-1482

N. parvipectoralis

~953

~6153

~603

—

N. daedalus

68-78

725-746

19-22

225-235

1 From Nelson 1966.

2 From Castle 1980.

3 Calculated from Chu, Wu, and Jin 1981.

McCOSKER: NEW WORM EELS

differences. In an independent study, Mark M.
Leiby (in litt, Florida Dept. of Natural Resources,
14 Jan. 1982) has compared the leptocephali of
species of those genera and concluded that they
are trenchantly different. The two Pseudomyro-
phis species that are similar in body elongation
to Neenchelys daedalus, P. nimius BOhlke, 1 960,
and P. atlanticus Blache, 1975, inhabit mud and
sand substrates in deep water. Dean (1972) and
Mark Leiby (in litt.) have concluded that Myro-
phisfrio Jordan and Davis, 1891, and an unde-
scribed Atlantic species are congeners of Pseu-
domyrophis nimius. Two species are known from
the eastern Pacific: P. micropinna Wade, 1946,
the type-species; and an undescribed species
ranging from Costa Rica to Baja California.

Eleven other Pacific specimens, captured by
midwater trawls, were tentatively identified as
Neenchelys daedalus. They are all smaller spec-
imens and appear identical in proportions to the
new species. They differ considerably, however,
in total vertebrae numbers: the holotype and
paratype have 235 and 225, respectively, whereas
eight of the others had 25 1-274 (Jc = 266.8) ver-
tebrae. I am unable to account for such a large
mean difference and broad range in vertebral
number for conspecifics in such close geograph-
ical proximity, and therefore have not made them
type-specimens.

COMPARATIVE MATERIAL.— Neenchelys daedalus (non-para-
types): AMS 1.19707-017 (field no. JP 69-53), 5(172-187),
Manus Island, Papua New Guinea, 4°1 5'S, 1 45°1 1 'E, 6-ft IKMT,
0-125 m, over 750+ m depth, John E. Paxton aboard FRY
TAGULA, 0120-0320 h, 22 Oct. 1969. (Many gonostomatids
and myctophids were captured in the same collection.) From
the same collection: CAS 50709, 2( 1 87-225); CAS 507 1 0, 1 ( 1 90,
cleared and stained); and ANSP 149295, 2(175-185). SIO 77-
171 1(144), Banda Sea, 105 km sw of Buru Is., 04°30.5'S,
125°34.6'E, 0-1500 m over 3600 m, Jim Coatsworth, 26 Aug.
1976.

Neenchelys buitendijki: ZM A. 102. 171, 1(218), syntype,
"probably from Moluccos," Indonesia. UH uncat., 2(1 18-123,
specimens dissected), Bombay City, India (specimens from
Mohamed 1958, reported on by Nelson 1966).

Neenchelys microtretus: BMNH 1 9 1 5. 1 0.25. 1 , 1 ( 1 83), holo-
type, Red Sea.

Pseudomyrophis micropinna: LACM 21557, 1(139), holo-
type, Isla Ladrones, Gulf of Chiriqui, Panama. SIO 60-72, head
and trunk only, cleared and stained.

Pseudomyrophis nimius: USNM 186274, 1(319), holotype,
Gulf of Mexico. ANSP 110150, 1(350, cleared and stained),
Gulf of Mexico.

Pseudomyrophis atlanticus: MNHN 1971-40, 1(259) holo-
type, Pointe-Noire. MNHN 1971-41, 1(241), paratype, Pointe-
Noire.

ACKNOWLEDGMENTS

Specimens and assistance were generously pro-
vided by the staffs of many institutions. In par-
ticular, I thank: John Paxton and Doug Hoese,
Australian Museum, Sydney (AMS); Eugenia
Bohlke and the late James Bohlke, Academy of
Natural Sciences of Philadelphia (ANSP); the staff
of the British Museum (Natural History)
(BMNH); John E. Randall, Bernice P. Bishop
Museum (BPBM); William N. Eschmeyer and
staff, California Academy of Sciences (CAS);
Robert Lavenberg, Natural History Museum of
Los Angeles County (LACM); Jacques Blache
and Marie-Louise Bauchot, Museum National
d'Histoire Naturelle, Paris (MNHN); Richard H.
Rosenblatt, Scripps Institution of Oceanography
(SIO); William Bussing, University of Costa Rica
(UCR); Leighton R. Taylor, Jr., Waikiki Aquar-
ium and University of Hawaii (UH); the staff of
the United States National Museum of Natural
History (USNM); and Hans Nijssen, Zoologisch
Museum Amsterdam (ZMA). I also thank: Beth
A. Meinhard for the preparation of Figures 1-3;
Kathy Smith for the preparation of Figures 4-5;
Michael Hearne for the preparation of radio-
graphs; Lillian Dempster for nomenclatural
assistance; and Mark Leiby, Peter Castle, and
Eugenia BShlke for their stimulating discussions
concerning the relationships of worm eels.

LITERATURE CITED

BAMBER, R. C. 1915. Reports on the marine biology of the
Sudanese Red Sea, from collections made by Cyril Cross-
land. XXII. The Fishes. J. Linn. Soc. London 31:477-485.

BLACHE, J. 1975. Contribution a la connaissancedesPoissons
Anguilliformes de la c6te occidentale d'Afrique. 1 5C note:
complements aux families des Muraenidae, des Heteren-
chelyidae et des Ophichthidae. Bull. Inst. Fr. Afr. Noire
37(3):708-740.

BLEEKER, P. 1853. Diagnostische beschrivnigen van nieuwe
of wenig bekende wisschsoorten van Batavia. Tintal I-VI,
Nat. Tijdschr. Neder.-Indie 4:451-516.

BOHLKE, J. E. 1960. A new ophichthid eel of the genus Pseu-
domyrophis from the Gulf of Mexico. Not. Nat. (Phila.) 329:
1-8.

CASTLE, P. H. J. 1972. The eel genus Benthenchelys (fam.
Ophichthidae) in the Indo-Pacific. Dana Rep. 82:1-32.

. 1 980. Larvae of the ophichthid eel genus Neenchelys

in the Indo-Pacific. Pac. Sci. 34(2): 165-171.

CHU, Y. T., H. Wu, AND X. JIN. 1981. Four new species of
the families Ophichthyidae and Neenchelidae. J. Fish. China
5(l):21-27.

COHEN, D. M., AND D. DEAN. 1970. Sexual maturity and
migratory behaviour of the tropical eel, Ahlia egmontis.
Nature 227(5254): 189-1 90.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 5

DEAN, D. M. 1972. Osteology and systematics of the echeline

worm eels of the Atlantic Ocean (Pisces, Anguilliformes).

Ph.D. Thesis. Univ. of Miami, Miami, Florida. 88 p.
FOWLER, H. W. 1934. Descriptions of new fishes obtained

1907 to 1910, chiefly in the Philippine Islands and adjacent

seas. Proc. Acad. Nat. Sci. Philadelphia 85:233-267.
GOSLINE, W. A. 1951. The osteology and classification of the

ophichthid eels of the Hawaiian Islands. Pac. Sci. 5(4):298-

320.
JORDAN, D. S., AND B. M. DAVIS. 1891. A preliminary review

of the apodal fishes or eels inhabiting the waters of America

and Europe. Rep. U.S. Comm. Fish. Fish. (1888) Pt. 16:

581-677.
LOTKEN, C. F. 1851. Nogle bemaerkinger om naeseborenes

. . . aalefamilien. Vidensk. Meddel. Naturhist. Foren. Kjo-

benjavn. 21 p.
McCosKER, J. E. 1 970. A review of the eel genera Leptenche-

lys and Muraenichthys, with the description of a new genus,

Schismorhynchus, and a new species, Muraenichthys chilen-
sis. Pac. Sci. 24(4): 506-5 16.

1 977. The osteology, classification and relationships

of the eel family Ophichthidae. Proc. Calif. Acad. Sci., ser.

4,41:1-123.
MOHAMED, K. H. 1 958. On the occurrence of the eel Neenche-

lys buitendijki Weber & deBeaufort in Indian waters. J. Bom-
bay Nat. Hist. Soc. 55:51 1-517.
NELSON, G. J. 1966. Osteology and relationships of the eel

Neenchelys buitendijki. Copeia 1966:321-324.
TAYLOR, W. R. 1967. An enzyme method of clearing and

staining small vertebrates. Proc. U.S. Natl. Mus. 122(3596):

1-17.
WADE, C. B. 1 946. Two new genera and five new species of

apodal fishes from the eastern Pacific. Allan Hancock Pac.

Exped. 9(7): 18 1-2 13.
WEBER, M., AND L. F. DEBEAUFORT. 1916. The fishes of the

Indo-Australian Archipelago. III. Ostariophysi: II Cypri-

noidea, Apodes, Synbranchi. E. J. Brill, Leiden. 455 p.

CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, California 94 1 1 8

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 6, pp. 67-76, 3 figs., 3 tables.

July 6, 1983

PSILORHYNCHUS GRACILIS, A NEW CYPRINOID FISH
FROM THE GANGETIC LOWLANDS

By
Walter J. Rainboth

Department of Biology, University of California, Los Angeles,
Los Angeles, California 90024

ABSTRACT: Psilorhynchus gracilis is described from 147 specimens collected at 10 lowland localities in
Bangladesh. Of the four previously described species of Psilorhynchus (balitora, homaloptera, pseudecheneis,
and sucatio), the new species resembles P. balitora most closely. The new species is compared with /'. balitora
and P. sucatio, both of which are syntopic species of the Gangetic plains. P. sucatio may be distinguished
from P. balitora and the new species by its reduced dorsal ray count and numerous shape and coloration
differences. P. balitora differs from the new species in lateral line and circumferential scale counts, number
of unbranched pectoral rays, coloration, and shape of the head and mouth parts. The preferred habitat and
distribution of the three lowland species are compared, and a key to the genus Psilorhynchus is provided.

INTRODUCTION

Fishes of the genus Psilorhynchus McClelland
are known to occur primarily in the Gangetic
drainage of southern Asia, where four described
(Menon 1974) and one undescribed species are
found. Most ichthyologists (following Hora 1925)
have recognized Psilorhynchus as the sole genus
in the family Psilorhynchidae. A recent re-anal-
ysis of the relationships of Psilorhynchus has been
published by Chen ( 1 980), who believes the genus
to belong to the Cyprinidae. I have not located
a copy of that publication and must reserve com-
ment at this time.

Of the five known species, two (Psilorhynchus
homaloptera Hora and Mukerji, 1925, and P.
pseudecheneis Menon and Datta, 1 964) have rel-
atively smaller scales and greater numbers of
simple pectoral rays than the remaining three
species. These small-scaled species also prefer
high-gradient streams located in the eastern
Nepalese Himalayas (P. pseudecheneis) and the

Naga Hills of the Assam-Burma border (P. hom-
aloptera). The subspecies P. homaloptera rowleyi
Hora and Misra, 1 94 1 , of the Chindwin River
(Irrawaddy) is one of the two species of this genus
from Burma (Fig. 1). These taxa from high-gra-
dient streams are not known to present any taxo-
nomic problems.

Two species from easily accessible lowland
areas were described by Hamilton in 1822 as
Cyprinus sucatio and C. balitora. While collect-
ing in the People's Republic of Bangladesh in
1977 and 1978, I obtained three species from
Gangetic lowland streams. Two of the species
were those described by Hamilton and one is
described, herein, as new.

Hamilton's (1822) somewhat brief original
descriptions of two lowland species now referred
to Psilorhynchus were not accompanied by fig-
ures, although he had prepared figures for them
during his stay in India. Hamilton's figures are
of prime importance because he kept no pre-

167]

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 6

FIGURE 1 . Distribution of the known species of Psilorhynchus. Inset map indicates region displayed. Darkened circles represent
P. gracilis localities, with triangle marking the type-locality. Open circles are localities for P. balitora, with question marks in
outlying non-verified reports. Two localities have both P. gracilis and P. balitora. Vertical hatching covers the range of P.
pseudecheneis. Diagonal hatching shows the range of P. homaloptera. Stippling highlights the known range of P. sucatio.

served material. However, only half of the figures
(and neither of the Psilorhynchus figures) were
published in Hamilton's original monograph. The
remaining figures were published subsequently
by several authors including McClelland (1839),
who published Hamilton's figures of Psilorhyn-
chus. The reasons for this were explained par-
tially by McClelland (1839) and more fully doc-
umented by Gudger (1924) and Hora (1929).
Although he had seen but a single individual of
one of the species, McClelland placed both species
in the new genus Psilorhynchus and presumed
that his single specimen was P. balitora (Ham-
ilton), for which he coined a new specific name,
variegatus, to replace balitora— a local term
meaning sand-digger. McClelland preferred the
use of classical Latin or Greek to the use of local
dialects; for example, he changed the homalop-
terid generic name Balitora Gray, 1834, to Pla-
tycara. Such replacement of names is inadmis-
sible under the present rules of nomenclature.

In Hamilton's figures (McClelland 1839:pl. 50),
Psilorhynchus sucatio is easily recognized; how-
ever, I collected two species which resemble the
illustration of P. balitora. Because McClelland's
P. variegatus refers to an individual of a balitora-
like species, the possibility that his species may

have been distinct from Hamilton's will also be
explored.

MATERIALS AND METHODS

Measurements were made to the nearest 0.1
mm with dial calipers. Standard length was mea-
sured from the snout tip to the base of the caudal
fin. All pre-fin lengths extend from the tip of the
snout to the base of the first unbranched ray at
fin origin. Dorsal and anal fin lengths are mea-
sures of the longest simple ray, whereas the paired
fins are depressed and measured from insertion
to posteriormost extension. Caudal fin length is
measured from the posterior end of the urocen-
trum to the tip of the normally spread upper fork.
Body depth is measured from dorsal fin origin
to pelvic fin insertion. The snout to occiput length
(head length = HL) is the distance to the poste-
rior margin of the supraoccipital bone. The snout
to preopercle length is taken on a horizontal.
Snout to maxilla length is measured to the pos-
terior end of the maxilla. Orbital measurements
are taken to the bony margin. Mandible length
is distance from the symphysis to articulation
with the quadrate. Gape width is the distance
between the two articulation points of upper and
lower jaw.

RAINBOTH: NEW PSILORHYNCHUS

Fin-ray counts are expressed with lower case
Roman numerals signifying unbranched rays and
Arabic numerals for branched rays. The deeply
divided final branched ray in both the dorsal and
anal fin is counted as one. Caudal counts list
procurrent rays of the upper fork as Roman
numerals with Arabic numerals for principal rays
of the upper/lower forks followed by Roman
numerals giving lower-fork procurrent rays. Lat-
eral line scales were counted for the body and
tallied separately from those on the caudal fin
base. Lateral transverse counts include the
median scale at the dorsal fin origin, record the
lateral line with a slash (/), and include the median
ventral row before the anal fin. Circumferential
counts encircle the body on the scale row imme-
diately anterior to the dorsal and pelvic fins. Cir-
cumpeduncular counts include all scales around
the peduncle at its narrowest region. The number
of anal scales refers to median scale rows between
the anus and anal fin. Belly scale rows include
all complete, free-edged midventral scale rows
crossing anterior to the pelvic fin insertion. Counts
given for lateral blotches include all distinct mid-
lateral blotches whether or not they are perfectly
bisected by the lateral line.

Body measurements are summarized as per-
cent standard length (% SL), head measurements
as percent head length (% HL).

Most of the material was collected by the author
and is housed at the Museum of Zoology, The
University of Michigan, Ann Arbor (UMMZ) or
at the Chandpur Freshwater Fisheries Research
Station, Chandpur, Bangladesh (CFRS). Other
specimens are from the American Museum of
Natural History, New York (AMNH), and the
Stanford University (SU) collection now housed
at the California Academy of Sciences, San Fran-
cisco. Paratypes of the new species have been
deposited at all aforementioned institutions plus
the Field Musuem of Natural History, Chicago
(FMNH), the Academy of Natural Sciences of
Philadelphia (ANSP), and the United States
National Museum, Washington, D.C. (USNM).

Distribution maps are part of drainage maps
of the southern half of the Asian continent drafted
by the author from the most recent world-wide
series of l:5,000,000-scale topographic maps
prepared by the U.S. Defense Mapping Agency.

Psilorhynchus McClelland

Psilorhynchus MCCLELLAND, 1839:300, 428 (type-species
Cyprinus sucatio Hamilton, by subsequent designation of
Jordan 1919).

DIAGNOSIS.— Body arched dorsally and flat-
tened ventrally. Anteriorly depressed, becoming
cylindrical with lateral compression posteriorly.
Ventral surface of head markedly flattened.
Mouth small, inferior, and transverse with a pro-
jecting snout; devoid of barbels. Pharyngeal teeth
uniserial. Gill membranes joined broadly to isth-
mus with aperture extending ventrally to base of
pectoral fin. Paired fins inserted horizontally.
Breast naked. Scales moderate to large, 31 to 50
pored scales on the complete lateral line. Fin-ray
counts: D ij-iii/7-9; A ii-iii/5; P, iv-x/9-12;
P2ii/7. Anus very close to pelvic fin; at least 8
scale rows separate anus from anal fin.

Key to the Species of Psilorhynchus

la. Branched dorsal fin rays 7 2

Ib. Branched dorsal rays 8 (or 9) 4

2a. Abdomen fully scaled; simple pectoral rays
4 sucatio (Hamilton)

2b. Abdomen naked; simple pectoral rays 7-
1 0 3

3a. Total lateral line scales 42-44; simple pec-
toral rays 7 or 8; abdomen smooth

homaloptera Hora and Mukerji

3b. Total lateral line scales 48-50; simple pec-
toral rays 10; abdomen with three trans-
verse folds of skin

pseudecheneis Menon and Datta

4a. Gape width greater than mandible length;
simple pectoral rays 6-7 (rarely 5) or more;

circumferential scales 1 8

balitora (Hamilton)

4b. Gape width much less than mandible
length; simple pectoral rays 4-5; circum-
ferential scales 16 gracilis sp.nov.

Psilorhynchus gracilis sp.nov.

(Figures 2 and 3)

HOLOTYPE.-UMMZ 205342 (adult female, 50.5 mm SL),
Jabuneswari River at Badarganj, Rangpur Dist., Bangladesh,
3 Apr. 1978.

PARATYPES (all from Bangladesh). -UMMZ 205343 (26
specimens, 29.4-49.9 mm SL), and CFRS uncat. (9 spec.) same
collection data as holotype. UMMZ 205337 (2 spec., 25.7-
29.7), Sangu River at Bandarban, Chittagong Hill Tracts, 25
Dec. 1977; UMMZ 205345 (8, 34.1-40.7), Ghaghat River at
Rangpur, Rangpur Dist., 3 Apr. 1978; UMMZ 205348 (31,
26.6-51.2), USNM 231693 (5), AMNH 43097 (5), CAS 5001 1
(5), and ANSP 148729 (5), and FMNH 94285 (5), Mahananda
River at Tetulia, Dinajpur Dist., 5 Apr. 1 978; UMMZ 20535 1
(15, 27.6-46.4), Keratoya River at Bhajanpur, Dinajpur Dist.,
6 Apr. 1978; UMMZ 205353 (2, 34.3-39.0), Tangam River
at Thakurgaon, Dinajpur Dist., 6 Apr. 1978.

OTHER MATERIAL EXAMINED (all from Bangladesh).— UMMZ

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 6

FIGURE 2. Young specimens of three lowland species of Psilorhynchus from Bangladesh, (top) P. gracilis, 34 mm SL,
Jabuneswari River, (middle) P. balitora, 20 mm SL, Mahananda River, (bottom) P. sucatio, 64 mm SL, Rangapani Creek.

205340 (18, 10.4-19.0), Dharla River at Kurigram, Rangpur
Dist., 2 Apr. 1978; CFRS uncat. (3, 33.8-35.4), Muhuri River,
10 km NE of Feni, Noakhali Dist., 2 Feb. 1978; CFRS uncat.
(1, 40), Halda River at Daulatpur, 40 km N of Chittagong,
Chittagong Dist., 24 Feb. 1978; CFRS uncat. (6) Halda Creek
at Khaia Bazaar, 53 km N of Chittagong, Chittagong Dist., 24
Feb. 1978.

DIAGNOSIS. — Lateral line scales 33 to 36 on
body plus 1 or 2 pored scales on caudal fin, to-

talling 35 to 37; pectoral fin with 4 or 5 simple
rays; branched dorsal rays 8 (rarely 9); 2 distinct
dorsal spots anterior to dorsal fin origin with a
third at the origin; midventral region scaleless
anteriorly, with 2 to 5 complete, free-edged scale
rows immediately anterior to pelvic fin insertion;
pectoral fin short, extending beneath dorsal fin
origin, but never reaching pelvic insertion; eye

RAINBOTH: NEW PSILORHYNCHUS

FIGURE 3. Comparison of ventral (A) and dorsal (B) aspects in Psilorhynchus balitora and P. gracilis. In each pair, left is P.
balitora, and right is P. gracilis. In ventral aspect (A), P. balitora, 39 mm SL, Siliguri, and P. gracilis, 47 mm SL, Jabuneswari
River. In dorsal aspect (B), P. balitora, 27 mm SL, Mahananda River, and P. gracilis, 32 mm SL, Jabuneswari River.

in upper part of head, not visible from below;
no spots on anal or paired fins, although mem-
brane between first 5 pectoral rays slightly dark-
ened in some adults.

DESCRIPTION.— Predorsal scales 10 or 11; cir-
cumferential scales 1 6 (rarely 1 5 or 1 7); circum-
peduncular scales 10; lateral transverse scale-rows
4/3; scale rows between anus and anal fin 8 or
9. Scales one row above lateral line immediately
posterior to dorsal fin have 6 or 7 radii in adults
of 50 mm SL.

Paired fins inserted horizontally; pectoral with
4 or 5 unbranched rays and 9-12 branched rays,
totalling 14-17 elements, extending as far as dor-
sal fin origin when depressed, with distal margin
separated from pelvic fin insertion by at least 2
scale rows; pelvic fin inserted slightly posterior
to dorsal fin origin, with 2 simple and 7 branched
rays. Dorsal fin closer to snout tip than to caudal
base, with 3 simple and 8 (rarely 9) branched
rays. Anterior and posterior rays of equal exten-
sion in depressed dorsal fin, which when nor-
mally expanded has an oblique and concave pos-
terior margin. Anal fin short and somewhat

falcate, its posterior margin well in advance of
caudal fin, with 3 simple and 5 branched rays.
Caudal fin deeply and evenly forked, possessing
19(1 0/9) principal rays preceded by 6 or 7 pro-
current rays above and 5 or 6 below. Body elon-
gate, arched above and generally flat below;
greatest depth at dorsal fin origin and greatest
width at pectoral fin insertion; shape somewhat
depressed anteriorly, gradually becoming cylin-
drical, then compressed posteriorly.

Head depressed, ventral mouth small and
transverse. Upper lip separate from upper jaw
by a deep groove and from snout by a shallow
groove. Ventral surface of snout separated from
lateral surfaces by deep rostral grooves. Upper
lip joined to lower lip at corner of mouth by a
fairly prominent flap of skin. Lower lip thick and
followed on chin by a number of large, globular
papillae which decrease in size posteriorly. Lower
jaw long, its length much greater than width of
gape. Gill membranes broadly joined to isthmus;
gill aperture extends dorsally from pectoral fin
base. Eye large, upper margin level with flat inter-
orbital space; width of orbit approximating inter-

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 6

orbital width. Tip of snout somewhat pointed if
viewed from dorsal aspect.

Nuptial tubercles evident on large males, ran-
domly distributed on head. Heaviest tubercle
accumulation below eye on lower part of cheek
where eruption takes form of tight clusters rather
than usual single pattern. Anterior body scales
have tubercles bordering their free margins.
Tubercle distribution on body progressively
sparser posteriorly. Pectoral fins on breeding
males show single row of tubercles ornamenting
dorsal surface of each ray, with tubercles on ante-
rior rays largest. Pelvic fins with same pattern
but with smaller tubercles. Large females have
same tubercle distribution as males but with slight
differences. Females have lower tubercle density
than males, lack tubercles on paired fins, and do
not display tubercles in the clumped suborbital
pattern of males.

COLORATION. — Generally recognizable in
specimens greater than 1 5 mm SL, although vari-
able. Laterally, a series of 7-10 dark blotches
with posteriormost extending onto caudal fin.
First lateral blotch midway between lateral line
and base of pectoral fin in small specimens. This
spot enlarges with age, to cover lateral line. Lat-
eral line always bisects remaining large lateral
spots. Middorsal spots discontinuous with lateral
spots; dorsal spots on same transverse scale rows
as lateral blotches, having identical counts on all
specimens. Where fish has unequal lateral spot
counts, some dorsal spots obliquely cross mid-
dorsal line, and on each side are still on same
scale rows as lateral spots. Two distinct dorsal
spots anterior to dorsal fin origin, with another
at origin and a fourth under posterior dorsal fin
rays. Four or five spots between dorsal fin and
caudal fin. All scales of back and upper side have
marginal melanophores connecting to create net-
work ventrally to just below lateral line. Network
darkens with age, but always distinct. Head dark
dorsally with light median-longitudinal streak
flecked with large discrete melanophores from
snout to posterior edge of occiput. A dark band
projects from each nostril to coalesce anteriorly
at tip of snout. Pigment lacking on fins of small
specimens, appearing gradually at 30 mm SL and
increasing substantially with size. Caudal fin ulti-
mately has 2 black blotches per lobe, with prox-
imal spot often joining median spot at caudal
base. Dorsal fin has apical spot which runs along
anterior 2 or 3 rays. No blotches on paired fins,

although large specimens have darkened mem-
branes between first 4 or 5 pectoral rays. Peri-
toneum dense black dorsally, becoming an irreg-
ular medium to dark gray below.

ETYMOLOGY. — The Latin adjective gracilis
(gender masculine) was chosen because of the
fish's slender shape, which immediately distin-
guishes the new species from Psilorhynchus bal-
itora, the most similar species.

COMPARISON OF SPECIES

Several counts aid in distinguishing the three
lowland species. The diagnostic counts (Table 1)
demonstrate resemblances between P. gracilis and
P. balitora in dorsal rays and total pectoral fin
elements. However, lateral line and circumfer-
ential counts are closest between P. gracilis and
P. sucatio. Belly squamation is distinctive in P.
gracilis, which never has more than two to five
midventral scale rows anterior to the pelvic fins.
P. sucatio always has a fully scaled belly and
sympatric specimens of P. balitora never exhibit
fewer than eight midventral prepelvic scale rows.
Specimens from the Chindwin River, Burma, and
the Rapti River, Nepal have naked breasts and
bellies, but are typical P. balitora in all other
respects. Both P. homaloptera and P. pseude-
cheneis have scaleless bellies. Two AMNH 15767
paratypes of P. homaloptera from the Naga Hills
were incorrectly identified. One is P. balitora with
typical counts and measurements which have
been included in all four tables. The other spec-
imen is a loach (genus Noemacheilus).

Several proportional measurements show
notable differences. However, overlap in pro-
portional measurements may occur when juve-
niles are included, even though adult proportions
display pronounced differences. For instance, of
seven measured P. balitora, four adults were over
40 mm SL and three juveniles were less than 30
mm SL. The robust adults have a body depth of
25.5% SL, whereas the juveniles have a body
depth of 19.4% SL (Table 2). Juvenile propor-
tions of P. balitora overlap with those of adults
of the other two species. Other proportions dis-
tinguishing P. gracilis from P. balitora at any size
are those for the anal, pectoral, and pelvic fins.
Proportions (taken as % HL) differing consis-
tently in these two species are interorbital width,
gape width, and mandible length (Table 2). A
single perfectly discriminating character is gape
width, which is considerably less than the man-

RAINBOTH: NEW PSILORHYNCHUS

TABLE 1. DIAGNOSTIC COUNTS FOR THE THREE LOWLAND SPECIES OF PSILORHYNCHUS. Number counted in parentheses.

P. balitora

P. gracilis

P. sucatio

Dorsal fin rays
Pectoral fin rays
Caudal fin rays
Lateral line scales
Circumferential scales
Anal scales

iii/8 (17)
v-viii/7-9 (41)
v-vi, 9/8, iv-v (10)
30-34+ 1 or 2 (41)
17-20(41)
9-10(17)

iii/8-9(18)
iv-v/9-12(60)
vi-vii, 10/9, v-vi (18)
33-36 + 1 or 2 (60)
15-17(60)
8-9 (18)

ii/7 (18)
iv/8-9 (18)
iii-iv, 9/9, hi (18)
34-35+ 1 or 2 (18)
16-18(18)
8-11 (13)

dible length in P. gracilis, whereas the size re-
lationship is reversed in P. balitora.

Psilorhynchus gracilis differs from P. sucatio
in the depth of the caudal peduncle and several
head-measurement proportions, most notably the
interorbital width: approximating the orbital
width in P. gracilis and nearly doubling the orbital
width in P. sucatio. Also, in P. sucatio the ante-
rior dorsal rays show greater extension when
depressed, whereas P. gracilis and P. balitora
have equal or greater posterior extension of the
last rays.

Color patterns are most similar between P.
gracilis and P. balitora in younger specimens.
Therefore, sub-adults have been illustrated (Fig.
2). Juveniles of P. sucatio have a continuous black
midlateral stripe extending from opercle to cau-
dal fin. The stripe fades with age, and gradual
coalescence of blotches gives the adult color pat-
tern. Remnants of the stripe can be seen on the
figured specimen as discrete melanophores

between the lateral spots. Dorsally, P. sucatio
exhibits darkening of entire scales rather than
the reticulated network found in P. gracilis and
P. balitora. Both P. gracilis and P. balitora always
have a series of lateral blotches with smaller and
more numerous spots on P. gracilis (Table 3).
The lateral spots on P. gracilis and P. balitora
are on the same diagonal scale rows as the dorsal
spots. On P. gracilis dorsal and lateral spots are
totally distinct, whereas P. balitora exhibits a
faint continuous band between the lateral and
dorsal spots. The predorsal spot pattern (Fig. 3)
for P. gracilis is two distinct blotches with a third
at the dorsal fin origin, and for P. balitora a single
blotch, with a second beginning at the dorsal
origin. In adult P. balitora the dorsum gradually
darkens causing the pattern to become somewhat
obscure, although it still persists. P. gracilis retains
a distinct reticulated pattern throughout adult
life.
A recently described taxon Psilorhynchus

TABLE 2. PROPORTIONAL MEASUREMENTS FOR THREE SPECIES OF PSILORHYNCHUS. Characters 2 through 9 expressed as %SL.
Characters 10 through 15 expressed as %HL (snout to occiput).

P. balitora

P. gracilis

P. sucatio

Range (mm)

X ± so

Range (mm)

X ± so

Range (mm)

Standard length
Snout to dorsal fin

22.7-47.8
48.1-52.8

(n
50.3

7)
2.1

28.2-51.2
44.6^7.9

(n
46.8

= 10)
± 1.1

26.4-64.3
43.9^18.9

(n =
46.3 :

= 13)
b 1.6

Snout to pectoral fin

22.4-23.9

23.2

0.5

18.8-22.7

20.1

± 1.3

19.7-23.1

21.0 :

b 1.1

Body depth

18.5-26.9

22.9

3.5

18.0-21.4

19.7

± 1.3

15.7-21.4

18.3 :

b 1.9

Peduncle depth

8.2-9.4

8.9

0.4

7.5-8.4

8.0

± 0.3

6.1-7.3

7.0 :

b 0.4

Pectoral fin length

25.9-28.7

27.4

1.0

21.0-23.4

22.4

± 0.8

17.3-22.7

20.6 :

b 1.6

Pelvic fin length

20.7-22.9

21.5

0.3

16.9-19.9

18.4

± 1.0

16.4-19.3

18.4 :

b 1.0

Anal fin height

15.4-18.0

17.0

0.9

14.2-15.2

14.8

± 0.4

12.1-14.4

13.0 :

b 0.9

Snout to occiput

23.0-24.8

24.1

0.6

20.3-24.0

22.0

± 1.3

20.4-23.9

21.5 :

b 1.0

10.

Snout to maxilla

32.3-35.6

33.6

1.3

23.6-31.0

27.8

± 2.4

25.5-32.2

31.0 :

b 0.9

11.

Orbit width

31.3-35.4

33.4

1.5

30.3-33.8

31.8

± 1.0

25.2-30.5

27.7

1.7

12.

Interorbital width

36.5^1.6

38.6

2.0

29.8-36.3

33.7

± 2.0

42.8-56.5

50.0

4.7

13.

Gape width

26.8-31.6

28.4

2.0

19.1-24.7

22.1

± 1.6

23.3-28.2

25.6

1.9

14.

Mandible length

19.6-23.6

21.4

1.5

29.4-42.5

33.9

± 4.1

25.0-32.7

27.3

2.3

15.

Head depth at pupil

46.4-55.3

51.0

3.1

39.4.^7.3

43.8

± 2.8

33.6^2.0

38.6

2.7

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 6

TABLE 3. DISTRIBUTION OF VALUES FOR CHARACTERS
DISTINGUISHING PSILORHYNCHUS GRACILIS AND P. BALITORA.
Counts for P. gracilis holotype are underlined.

Total lateral line scales

P. gracilis

P. balitora

Simple pectoral rays

P. gracilis

P. balitora

Circumferential scales

P. gracilis

P. balitora

Lateral blotches

P. gracilis

P. balitora

sucatio var. damodarai David, 1953, appears to
be a local race of P. sucatio with slightly larger
pectoral fins than those found on individuals from
Bangladesh. The Damodar River variety is also
listed as having "8" branched dorsal rays as
opposed to "7-8" for Gangetic specimens. I have
not seen any specimens with eight branched dor-
sal rays, although the deeply divided last ray could
be erroneously counted as such.

DISCUSSION

Of the characteristics given for identification
in Hamilton's description (1822), few would be
of use in distinguishing species as similar as P.
gracilis and P. balitora. However, Hamilton does
state that there are approximately 1 2 rays in each
pectoral fin. This count is closer to what I found
for P. balitora (minimum 1 3) and fewer than in
the new species (Table 3).

Hamilton's figure of the dorsal aspect repro-
duced in McClelland (1839) shows six simple
rays in each pectoral fin, and anterior-dorsum
and head color patterns identical to my obser-
vations for P. balitora. Hamilton's figure dis-
plays the anterior part of the dorsum on P. bal-
itora with one blotch midway between the occiput
and the dorsal fin, and another at the dorsal fin
origin on both lateral and dorsal views. The dor-
sal coloration of the head has two black spots
separated by a transverse white line in the inter-
orbital space. There is no longitudinal white streak
from the snout to occiput as on the new species

(Fig. 3). There are 34 total scale rows in longi-
tudinal series, which better describes P. balitora
(Table 3). Although the lateral blotches on Ham-
ilton's illustration of the lateral aspect might cor-
respond to the new species, six lateral spots are
commonly seen in P. balitora, which also exhibits
a lengthened anterior blotch similar to that in
Hamilton's figure. The total dorsal spots illus-
trated in dorsal aspect might be high for P. bal-
itora, however, even though both lateral and dor-
sal views are presumably taken from the same
fish, the blotches do not match (the lateral view
has one fewer dorsal spot on the peduncle).
Because the lateral view shows an equal number
of lateral and dorsal blotches, that would pre-
sumably be a better indicator of the dorsal spot
pattern on the caudal peduncle than the illus-
trated dorsal aspect. From these characters it
would seem that Hamilton's Cyprinus balitora
is synonymous with my Psilorhynchus balitora
rather than with the new species.

Psilorhynchus variegatus McClelland remains
as a potential name for the new species.
McClelland (1839:430) stated that his single
specimen differed little from Hamilton's balitora
except for a few features. McClelland listed 17
rays in the pectoral fins and 33 scales in the lateral
line. The total of 1 7 pectoral fin elements occurs
in both species as a maximum, but 33 lateral line
scales is the mode for P. balitora and two scales
below the minimum total count found in P. gra-
cilis (Table 3). Therefore, it appears that P. var-
iegatus McClelland is indeed a synonym of P.
balitora (Hamilton) and that P. gracilis is a new
species.

Other accounts in Day (1878) and Shaw and
Shebbeare (1937) apply to P. balitora. No pub-
lished account appears to have included the new
species under the name P. balitora, which is rather
surprising in view of its abundance and appar-
ently widespread occurrence in the Gangetic low-
lands.

DISTRIBUTION AND HABITAT
PREFERENCE

Thus far the new species has been collected
only in Bangladesh. However, the extent of its
preferred habitat would suggest a much wider
distribution throughout the lower reaches of the
Ganges and Brahmaputra rivers. The southern-
most collections of Psilorhynchus gracilis in
Bangladesh are from rivers that currently have

RAINBOTH: NEW PSILORHYNCHUS

independent exits into the Bay of Bengal (Fig. 1).
The species is also fairly common in sandy
streams of northwest Bangladesh. Psilorhynchus
gracilis was taken in the same collections with
P. balitora and P. sucatio in the Mahananda River
at Tetulia and the Keratoya River at Bhajanpur,
both in Dinajpur District. P. sucatio was taken
at each of my collection localities for P. gracilis
except one site from the Muhuri River (Feni River
drainage) of Noakhali District in southeast Bang-
ladesh. However, P. sucatio was taken in two
other collections from the Feni River drainage.

Psilorhynchus gracilis is found over small peb-
bles in shallow running waters where the bottom
is primarily sand. In this regard it resembles P.
balitora which is sometimes found in the same
habitat but which is always closely associated
with hard substrates. P. gracilis is generally free-
swimming and occasionally rests on its spread
paired fins. P. balitora prefers to maintain close
fin contact with the substrate, often not moving
unless strongly disturbed. Psilorhynchus gracilis
may be caught easily on pebble outcrops having
both species present because of its greater ten-
dency to leave the bottom. A seine pulled under
or through the gravel of the same outcrop catches
P. balitora, which will be taken out along with
the substratum. P. sucatio differs from the other
lowland species in being taken primarily along
the edges of sandy streams, and seems to be most
abundant near emergent or overhanging vege-
tation. I have not observed Psilorhynchus bur-
rowing, although several species of loaches were
seen to burrow into sand within inches of Psi-
lorhynchus under observation. Attempts to elicit
burrowing by disturbing the fishes were not suc-
cessful.

The P. balitora taken at the same localities as
P. gracilis were all small, about half to two-thirds
adult size. This possibly indicates that fully grown
individuals occur upstream in areas of higher
gradient. It is also consistent with the physical
and behavioral characteristics of the species. P.
balitora has much larger pectoral fins with more
simple rays, and a wider and higher body pre-
dorsally. It depresses its head when positioned
in an area of strong current, and is forced down
onto its fins. This shape is common among Asian
hillstream fishes which attach themselves to hard
substrata in high-gradient streams (e.g., Garra,
Homaloptera, Gastromyzon). This somewhat
passive posturing for increased friction is a much

more efficient method than constant swimming
for maintaining position in the current of tor-
rential streams. Thus, it would appear that north-
ern Bangladesh may be the southernmost region
of the Gangetic plain occupied by P. balitora, a
species adapted to higher gradients than either
P. gracilis or P. sucatio,

COMPARATIVE MATERIAL EXAMINED

Psilorhynchus balitora— INDIA: SU 28701 (2 specimens, 42.8-
45.2 mm SL), Siliguri, North Bengal, no date given; SU 32627
(1, 40.2) Siliguri, North Bengal, Apr. 1937; AMNH 15767 (1,
47.8), Keleki Stream at Emilioni, Naga Hills, Assam (paratype
of Psilorhynchus homaloptera Hora and Mukerji). NEPAL:
UMMZ 207678 (7, 33.1-37.2), Rapti River at Chitawan Val-
ley, Apr.-May 1975. BURMA: AMNH 13811 (14, 32.5-42.8),
Upper Burma, Chindwin drainage. BANGLADESH: UMMZ
205347 (19, 15.6-26.7) Mahananda River at Tetulia, Dinajpur
Dist., 5 Apr. 1978; CFRS uncat. (15) same data; UMMZ 205350
(11, 14. 1-18.6), Keratoya River at Bhajanpur, Dinajpur Dist.,
6 Apr. 1978.

Psilorhynchus sucatio— BANGLADESH: UMMZ 205338 (5
specimens, 20.2-22.6 mm SL), Sangu River at Bandarban,
Chittagong Hill Tracts, 25 Dec. 1977; UMMZ 205339 (32,
46.3-73.3), Rangapani Creek, 6 km NNW of Jaintapur, Sylhet
Dist, 19 Feb. 1978; UMMZ 205341 (5, 12.5-17.5) Dharla
River at Kurigram, Rangpur Dist., 2 Apr. 1 978; UMMZ 205344
(5, 1 5.5-55.6), Jabuneswari River at Badarganj, Rangpur Dist.,
3 Apr. 1978; UMMZ 205346 (57, 13.8-58.8), Ghaghat River
at Rangpur, Rangpur Dist., 3 Apr. 1978; UMMZ 205349 (42,
15.7-41.8), Mahananda River at Tetulia, Dinajpur Dist., 5
Apr. 1978; UMMZ 205352 (22, 11.1-50.7), Keratoya River
at Bhajanpur, Dinajpur Dist., 6 Apr. 1978; UMMZ 205354
(37, 12.7-45.7), USNM 231694 (5), AMNH 43096 (5), and
50010 (5), ANSP 148728 (5), and FMNH 94284 (5), Tangam
River at Thakurgaon, 6 Apr. 1973; CFRS uncat. (19, 34.2-
71.9), Koilla Creek, 13 km w of Ramgarh, Chittagong Dist.,
3 Feb. 1978; CFRS uncat. (8, 25^*0), Feni River at Ramgarh,
Chittagong Hill Tracts, 3 Feb. 1978; CFRS uncat. (3, 46.4-
62.5), Tangam River at Kestapur, Dinajpur Dist., no date;
AMNH 19648 (2) Sevoke River, Darjeeling.

ACKNOWLEDGMENTS

My sincere thanks and appreciation go to Karl
F. Lagler, The University of Michigan, and John
R. Snell, A. Q. Chowdhury, and Kermit E. Sneed,
Irrigation Fishery Development Project, Bang-
ladesh, for help in obtaining and shipping the
specimens to The University of Michigan. I owe
a great deal to my close friend and associate Ataur
Rahman and many others in the Directorate of
Fisheries, Government of Bangladesh, who
greatly facilitated the field work. William N.
Eschmeyer (CAS) and Donn E. Rosen (AMNH)
loaned specimens. Reeve M. Bailey, Gerald R.
Smith, Barry Chernoff, and Michael L. Smith of
The University of Michigan offered helpful sug-

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 6

gestions pertaining to many aspects of manu-
script preparation. Tyson Roberts kindly donated
several specimens of P. balitora from Nepal to
the UMMZ. Mark Orsen and Kama Steelquist
of The University of Michigan gave much-
needed advice on illustrations. This work was
supported in part by the UCLA Department of
Biology Fisheries Program.

LITERATURE CITED

CHEN, Y. 1980. Acta Hydrobiologica Sinica (not seen).

DAVID, A. 1953. On some new records of fish from the Damo-
dar and Mahanadi river systems. J. Zool. Soc. India 5(2):
243-254.

DAY, F. E. 1878. The fishes of India, etc. 2 vols. London.

GRAY, J. S. 1 834. Illustrations of Indian zoology of new and
hitherto unfigured Indian animals from the collection of
General Hardwicke. 2 vols. London.

GUDGER, E. W. 1924. The sources of the material for Ham-
ilton-Buchanan's fishes of the Ganges, the fate of his collec-
tions, drawings and notes, and the use made of his data. J.
Proc. Asiatic Soc. Bengal 19(4): 12 1-1 36.

HAMILTON (formerly BUCHANAN), F. 1 822. An account of the

fishes found in the river Ganges and its branches. Edinburgh.

405 p., 39 pis.
HORA, S. L. 1925. Notes on fishes in the Indian Museum.

XII— The systematic position of the genus Psilorhynchus

McClelland. Rec. Indian Mus. (Calcutta) 27:457-460.
. 1929. An aid to the study of Hamilton Buchanan's

"Gangetic Fishes." Mem. Indian Mus. (Calcutta) 9(4): 1 69-

192.
, AND K. S. MISRA. 1941. Fishes collected by the

Vernay-Hopwood upper Chindwin Expedition, 1935. J.

Bombay Nat. Hist. Soc. 42(3):478^82.
, AND D. D. MUKERJI. 1935. Fishes of the Naga Hills,

Assam. Rec. Indian Mus. (Calcutta) 37:381^04.

MCCLELLAND, J. 1839. Indian Cyprinidae. Asiat. Res. 19:
217^171.

MENON, A. G. K. 1974. A check-list of fishes of the Hima-
layas and the Indo-Gangetic plains. Inland Fisheries Society
of India, Spec. Publ. 1.

, AND A. K. DATTA. 1961. Zoological results of the

Indian Cho-Oyu Expedition (1958) in Nepal. 7— Pisces. Psi-
lorhynchus pseudecheneis a new cyprinid fish from Nepal.
Rec. Indian Mus. (Calcutta) 59:253-255.

SHAW, G. E., AND E. O. SHEBBEARE. 1937. The fishes of
Northern Bengal. J. Asiat. Soc. Bengal, Science 3:1-137.

CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, California 94118

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 7, pp. 77-85, 1 fig., 5 tables.

July 6, 1983

SIZE AND DISTRIBUTION OF THE CALIFORNIA
SEA LION POPULATION IN MEXICO

Burney J. Le Boeuf,{ David Aurioles,2 Richard Condit,1

Claudio Fox,2 Robert Gisiner,3 Rigoberto Romero,2

and Francisco Sinsel2

ABSTRACT: California sea lions were censused during the 1979 and 1981 breeding seasons on rookeries and
hauling grounds in the Gulf of California and the Pacific coast of Mexico. Correcting for underestimate bias
and uncensused islands yielded estimates of approximately 20,000 sea lions in the Gulf with an annual
production of 7,000 pups, and 63,000 sea lions on the Pacific coast with an annual production of 29,000 pups.
Counts on seven major rookeries in the Gulf in 1979 were 35% higher than in 1966. Sixteen percent of the
California sea lion population in the United States and Mexico (estimated at 145,000 animals) resides in the
Gulf and 46% inhabits the Pacific coast of Mexico.

INTRODUCTION

The California sea lion, Zalophus californi-
anus, along with numerous species of birds and
whales, is a top trophic-level predator in the pro-
ductive waters of the Gulf of California. In the
southern part of the Gulf, it feeds predominantly
on a variety of fishes and occasionally on squid
(Aurioles, Fox, and Sinsel 1981). The full impact
of this pinniped on Gulf of California fisheries
is unknown because few systematic censuses of
these animals have been conducted.

Long before California sea lions in the Gulf of
California were counted, they were exploited.
Early inhabitants of Baja California, Sonora, and
Sinaloa killed California sea lions for their meat.

1 Department of Biology and Center for Coastal Marine
Studies, University of California, Santa Cruz, California 95064.

2 Centre de Investigaciones, Biologicas de Baja California,
La Paz, Baja California, Mexico.

3 571 West B Street, Dixon, California 95620.

Between 1860 and 1870, whalers killed thou-
sands of sea lions for their oil and skins. Later,
the animals were hunted for their "trimmings"—
vibrissae and genitalia which were sold in the
Orient as aphrodisiacs. Local slaughter for oil
and meat continued until the early 1960's on
some islands in the Gulf, e.g., Puerto Refugio on
Angel de la Guarda (Lluch 1969), and poaching
is still going on today.

The first systematic census of the sea lions in
the Gulf was conducted by Lluch ( 1 969). He esti-
mated 6,027 total animals on eight islands from
counts made in 1 963, 1 964, and 1 966 at the peak
of the breeding season. Orr, Schonewald, and
Kenyon (1970) censused several islands between
1960 and 1968, but with the exception of Los
Islotes, no rookery was censused during the
breeding season. Mate (1977) estimated 9,428
sea lions in the Gulf of California from aerial
censuses taken in June 1975, but his data are
difficult to compare with other censuses because

[77]

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 7

FIGURE 1 . The route of R/V Ellen B. Scripps on expedi-
tions in 1979 and 1981 from San Diego, California, to Cali-
fornia sea lion rookeries along the Pacific coast of Baja Cali-
fornia, Mexico, and in the Gulf of California. The itineraries
were similar in both years except for deviations taken in 1979
which are noted by solid lines. The numbers refer to rook-
eries not censused (1-6) and islands supporting nonbreeding
aggregations (7-12). The code is: 1 = Los Coronados; 2 = Isla
San Ger6nimo; 3 = Punta Rosarito; 4 = Isla Magdalena; 5 =
Isla Lobos; 6 = Farall6n at Mazatlan; 7 = Isla Salsipuedes; 8 =
Cabo Haro; 9 = Roca Vela; 10 = Isla Patos; 1 1 = Cabo Lobos;
1 2 = Punta Lobos.

they are categorized by latitude rather than by
island.

The main purpose of this report is to present
recent data on the number and distribution of
California sea lions in the Gulf of California dur-
ing the breeding season and to compare these
data to earlier counts and to counts obtained on
islands on the west coast of Baja California, Mex-
ico. In addition, we report weights and mea-
surements of sea lion pups obtained from Gulf
and Pacific rookeries.

METHODS

CENSUSES.— The data in this report were col-
lected on two expeditions aboard the Scripps
Institution of Oceanography research vessel Ellen
B. Scripps. Inclusive dates were 21 June to 21
July 1979 and 1 to 30 June 1981. The track
records are shown in Figure 1 .

Both expeditions were timed to coincide with
the breeding season of California sea lions and
to facilitate counting sea lions in the Gulf. At
Los Islotes, the peak number of breeding females
occurs at the end of June, and the maximum
number of pups are observed on about 10 July
(Auricles, Romero, and Fox 1978). We were aided
in locating all suspected rookeries in the Gulf by
previous investigations and by information
obtained from local fisherman. The only rook-
eries not censused in the Gulf were Isla Lobos
and the farallon at Mazatlan. We did not census
the following rookeries on the outer Pacific coast
of the peninsula: Isla Natividad, Los Coronados,
Isla San Geronimo,4 and Punta Rosarito. Esti-
mates of the sea lion population on these Gulf
and Pacific rookeries are available from other
sources (e.g., Mate 1977).

Censuses were conducted from a dinghy or
launch, from the ship, and on foot. Most counts
were made from a 4-m Avon or Zodiac inflatable
dinghy powered by a 15- or 25-hp outboard
motor. One person operated the boat and one or
two people censused. Small islands were circum-
navigated; on large islands, a dinghy was launched
after sea lion aggregations were sighted from the
ship. In ideal conditions, the boat was run slowly
within 10m of the shoreline. This did not alarm
the sea lions and accurate counts were obtained.
When waves or wind were strong, or extensive
kelp beds surrounded an island, the boat was
operated 30 to 70 m offshore. In bad weather,
binoculars were used from the research vessel
from 0.5 to 1 km offshore. This was necessary
at Isla Santa Margarita and three small non-
breeding aggregations on Isla Magdalena, Salsi-
puedes, and Cabo Haro in 1981.

We censused sea lions in six categories: adult
males, subadult males, females, juveniles, pups,
and miscellaneous. Adult males are the largest
animals with large dorsal head crests and thick
necks. Subadult males are smaller, have thinner
necks, and little crest development. For most
analyses, these two age categories are combined.
Females and juveniles are distinguished from
males by their smaller size, thinner neck, and
light brown color compared to dark brown in
males. Juveniles are the smallest class in this
group and include one- to two-year-olds, and
possibly some three-year-olds. The miscella-

1 Also spelled Isla San Jer6nimo.

CALIFORNIA SEA LION POPULATION IN MEXICO

neous category encompasses any animal except
pups that could not be readily categorized because
of high animal density or poor censusing con-
ditions. Pups are the unmistakable, but easily
concealed, young of the year.

VALIDITY AND RELIABILITY ESTIMATES.—
Validity tests were performed using ground counts
and boat counts of the same rookery. After a
census was conducted from a dinghy, one person
went ashore, counted adults from a hiding place,
and then frightened the adults into the water and
quickly counted the pups. This was done at six
places in 1 98 1 . In addition, at two sites we com-
pared counts from the ship with counts from a
dinghy. Estimates of reliability were obtained by
having two censusers make simultaneous counts
at all rookeries in 1979 and at five rookeries in
1981. For these areas, the mean of the two cen-
suses conducted is presented.

WEIGHTS AND MEASUREMENTS.— A crew of
three to six people weighed and measured 1 1 1
pups on six rookeries, one in the Pacific and five
in the Gulf. Landing near a rookery caused adults
to flee into the water leaving their pups behind.
The procedure consisted of capturing a one- to
three-week-old pup by the hindflippers and
restraining it on a flat rock. Curvilinear length
(tip of nose to tip of tail) was measured along the
contour of the dorsal midline. The pup was then
placed in a light leather dog harness and weighed
with a hand-held spring scale (2 5 -kg capacity ±
0.5 kg). After extracting a blood sample, the pup
was tagged with an "All-Flex" cattle ear tag in
the trailing edge of a foreflipper, its sex was deter-
mined, and it was carried back to where it was
captured. Mothers retrieved their pups shortly
after pups were released or soon after the launch
and crew departed from the rookery.

RESULTS

WEATHER.— Weather conditions in 1979 were
ideal for censusing in the Gulf; seas were calm;
winds of 0-5 knots were variable in direction.
Conditions were fair for censusing Pacific coast
islands except at Isla Santa Margarita where high
surf prevailed. Wind and choppy water made
censusing in the Gulf difficult in 1981, especially
on the windward sides of Islas San Esteban, San
Pedro Martir, Granito, and San Ignacio Farallon;
on the Pacific side, heavy surf made landings or
censusing by dinghy impossible at Islas Mag-

dalena and Santa Margarita and on the west side
of Isla Cedros.

GULF OF CALIFORNIA. — More than 15,000 sea
lions were counted in the Gulf in 1979 and more
than 14,000 in 1981 (Table 1). Major rookeries
in the Gulf were found on islands in the center
or midriff area: San Esteban and Angel de la
Guarda (Los Machos, Granito, and Los Can-
tiles). In 1979 and 1981, 76 and 71% of the sea
lions counted were observed in the area from the
north end of Angel de la Guarda (29°35'N) south
to San Pedro Nolasco (27°58'N). Most of the sea
lions in the north end of the Gulf were located
on Isla San Jorge. Relatively few sea lions breed
in the southern end of the Gulf, and these were
concentrated on Los Islotes and San Ignacio Far-
allon.

No pups were observed in the small groups of
sea lions present on Isla Salsipuedes, Cabo Haro,
Roca Vela, Isla Patos, Cabo Lobos, and Punta
Lobos. No sea lions were observed during careful
surveys of Isla Coronado, Tortuga, Raza, Mejia,
Cabo Tepopa, Los Frailes, Cabo San Lucas, and
the eastern half of Islas San Lorenzo, Las Ani-
mas, and Espiritu Santo. A few males or juveniles
were observed on Islas Carmen, San Lorenzo,
and Partida.

Eleven Gulf rookeries produced a minimum
of 3,422 pups in 1979, 2,277 in 1981 (Table 1).
The largest category of the animals counted on
rookeries were female (48% in 1979 and 60% in
1981). The mean operational sex ratio (adult
males:adult females) for rookeries was 1:8.30
(N = 9, so = 5.61) in 1979 and 1:9.41 (N = 10,
so = 3.83) in 1981. The overall operational sex
ratio for rookeries in these years were 1:5.84 and
1:8.94, respectively.

Sea lions in the Gulf occupied cobblestone
coves or the boulder- strewn shoreline. Few
breeding groups were seen on sand or gravel
beaches, which tended to be occupied by non-
breeding males. Most territorial males were
observed "patrolling" the water in front of the
females. Females and pups occupied a narrow
zone near the water's edge and rarely were
observed 10 m or or more inland. In some loca-
tions, rafts of females floated in the shallows and
the females occasionally vocalized to their pups
onshore. Even on islands supporting numerous
sea lions, the population was broken up into small
groups, and a small number of animals took up
a large part of the shoreline.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 7

TABLE 1. CENSUSES OF CALIFORNIA SEA LIONS IN THE GULF OF CALIFORNIA, 29 JUNE TO 8 JULY 1979 (top figures) AND 8-21
JUNE 1981 (bottom figures, in italics). "-" denotes that no census was taken. Abbreviations for census methods: G = ground,
L = launch, S = ship.

Census

Islands

methods

Males

Females

Pups

Juv.

Misc.

Totals

Los Islotes

G, L

138

San Ignacio Farall6n

—

G,L

133

323

Cabo Haro

San Pedro Nolasco

382

232

133

846

696

725

201

7,726

San Pedro Martir

150

903

321

167

1,634

G,L

134

1,061

252

720

725

7,692

San Esteban

G, L

504

2,044

820

325

268

3,961

G, L, S

295

2.252

705

239

702

3,593

Salsipuedes

Roca Blanca

121

207

Patos

454

473

Angel de la Guarda

Los Machos

709

168

404

1,316

L, S

762

226

247

1,389

Granito

436

337

242

1,102

G,L

658

161

1,018

Los Cantiles

163

969

446

219

1,810

688

270

702

229

1,380

Roca Vela

Cabo Lobos

170

San Jorge

167

1,398

1,030

632

3,253

765

2,034

457

667

3,344

Rocas Consag

—

295

421

Totals 1979

1,724

7,214

3,422

2,357

423

15,140

Totals 1981

988

8,639

2,277

1,789

696

14,389

Approximately 10-20% of the females
observed were still nursing yearlings, and some
small groups were made up almost entirely of
such pairs.

PACIFIC COAST.— Approximately 23,000 sea
lions were counted on Pacific coast rookeries in
1979, a greater number than in 1981 (Table 2).
However, some islands censused in 1979 were
not censused in 1981, and vice versa. A better
estimate of the total number of animals associ-
ated with the rookeries shown in Table 2 is
27,895— obtained by adding the 1981 census
count for Islas Cedros (4,730) and Magdalena
(47) to the 1979 total census.

At least 6,529 pups were produced on the

islands censused in 1979. If the number of pups
counted on Isla Cedros in 1981 (2,138) is added
to that, annual pup production was at least 8,667
pups.

As in the Gulf, females were in the majority
(53% of animals counted in 1979 and 55% in
1981). The mean operational sex ratio of rook-
eries in 1979 was 1:6.87 (N = 6, SD = 5.83), the
overall sex ratio was 1:3.58. Less than 2% of the
females were observed nursing yearlings.

Large aggregations of sea lions were observed
on each rookery visited with the exception of Isla
de Guadalupe. The females were hauled out in
clumps well above the surf line, usually on sandy
beaches. Territorial males stationed themselves

CALIFORNIA SEA LION POPULATION IN MEXICO

TABLE 2. CENSUSES OF CALIFORNIA SEA LIONS ON THE PACIFIC COAST OF BAJA CALIFORNIA, 13-20 JULY 1979 (top figures)
AND 3-5 JUNE 1981 (bottom figures, in italics). Numbers in parentheses are calculated from total number of animals censused;
"-" denotes that no census was taken. Abbreviations for census methods: G = ground, L = launch, S = ship.

Census

Islands

method

Males

Females

Pups

Juv.

Misc. Totals

San Martin

Guadalupe

L,S

100

San Benito

Oeste

G, L

355

502

283

1,153

G,L

775

750 287

Centra

G, L

817

3,718

2,560

7,183

1,985

7,755

3,287

Este

183

1,070

900

2,212

(97)

(441)

(304)

(JO)

852

Cedros

—

— —

G,L

270

1,880

2,138

282

760 4.730

Asuncion

G, L

1,793

4,183

1,582

506

8,064

(105)

(2.780)

(406)

(608)

3,899

Magdalena

—

— —

47 47

Santa Margarita

L, G

265

2,676

1,202

114

4,257

(108)

(1,086)

(652)

(46)

7,592

Punta Lobos

Totals 1979
Totals 1981

3,449
680

12,278
8,287

6,529
4,686

842
984

23,098
557 14.994

on land with the females. Nonbreeding males
were observed considerable distances inland on
nearby rocks or on unoccupied beaches. In smaller
groups that were sometimes seen in coves or on
rocky ledges, territorial males were either on land
or in the water near the females.

Eighteen sea lions were observed at sea in the
Pacific and in the Gulf in 1981. They were sol-
itary and most were within a few kilometers of
a rookery. Three exceptions were one sea lion
seen near Cabo Pulmo, one east of Cabo San
Lucas, and one near Isla Coronados.

CENSUS METHOD BIAS.— Comparison counts
using different census methods are shown in Table
3. Ground counts of pups exceeded those taken
from a launch, with the underestimate from a
launch being greatest when the animals were dis-
tributed in rocky areas. Launch and ground counts
of adults were similar; the slightly higher counts
from a launch were partly because launch counts
always preceded ground counts. The latter were
conducted in the morning when the temperature
was rising and some adults were entering the
water to cool off. Ship counts of adults were lower

than those taken from the launch, and pups were
most difficult to see from the large vessel.

Two measures of inter-observer agreement
were obtained. In 1979, independent counts of
two censuses on 13 rookeries were correlated.
Correlation coefficients were high and positive
for total animals (0.976), females (0.935), and
pups (0.926). Correlation coefficients were lower
for juveniles (0.777) and males (0.712), animals
which are more difficult to categorize, i.e., sub-
adult males or juveniles can be confused with
adult females. In 1981, two independent cen-
suses were conducted on parts of five rookeries
in the Gulf. The results, shown in Table 4, are
similar to those obtained in 1979 in that agree-
ment is usually high for total animals and females
and relatively low for juveniles. Independent
counts of pups in 1981 were more discrepant
than in 1979.

PUPS WEIGHTS AND MEASUREMENTS. — The
mean weight and curvilinear length of newborn
pups is shown in Table 5. On each rookery, the
mean weight and mean length of males is greater
than that of females. For all rookeries combined,

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 7

TABLE 3. A COMPARISON OF CENSUS METHODS ON SHORT SECTIONS OF ROOKERIES. Abbreviations for census methods: G
ground, L = launch, S = ship.

Island

Weather and terrain

Census
method

Adults

Pups

Cedros

Calm sea; animals packed tightly on sandy beach with

—

some in arroyo behind beach.

San Pedro Martir

Calm sea, overcast sky; animals on pebbly beach.

—

Granito (A.G.)

Calm sea on protected side; animals distributed on

269

small cobblestones & large boulders backed by ver-

231

117

tical cliffs.

San Ignacio

Calm sea; animals on rocky shoreline containing a

Farall6n

large open cave, flat shelves & large boulders.

San Esteban

Slight chop; animals on long sandy spit, in rock stub-

264

ble at base of vertical cliffs & in shallow sea caves.

388

Los Machos (A.G.)

Calm sea; animals on sandy beaches, rock tables &

347

among boulders at base of cliffs.

488

179

males were significantly heavier (t = 5.42, df=
109, P = < 0.05) and longer (/ = 4.88, df= 109,
P = <0.05) than females. The mean weights of
pups of both sexes from Cedros Island, the only
Pacific rookery represented, are lower than that
of pups from any Gulf rookery, but these differ-
ences are not statistically significant.

DISCUSSION

We counted 15,140 California sea lions in the
Gulf of California in 1979. This number is 61%
higher than an aerial census in July 1 975 by Mate
(1977) and 1 5 1 % higher than a launch and ground
census in 1963-1966 by Lluch (1969). Is our
higher count due to increased censusing effort,
the census methods employed, or to a genuine
increase in the population? Each explanation has
some validity.

Our census was more complete than Lluch's.
Fifty-four percent of the animals we counted in
1979 were on rookeries and resting places Lluch
did not visit; Los Islotes, San Esteban, Roca
Blanca, Angel de la Guarda (Los Machos and
Los Cantiles), Cabo Haro, Islas Salsipuedes and
Patos, and Cabo Lobos. If sea lions were found
on these islands in the mid-1960's in the same
relative proportions as in 1 979, the total number
of animals would have approximated 13,000,
16% fewer than we counted in 1979.

Terrain and climatic conditions in the Gulf are
such that an aerial census like that conducted by
Mate (1977) from a small airplane underesti-
mates the number of animals present more than
counts from a launch or on foot. Mate explains

that his count of 9,428 was low because of the
difficulty of counting and photographing animals
from the air over the dark, rocky terrain, with
many animals in the water by mid-morning. An
indication of the different results of these two
censusing methods is that Mate counted only 1 22
pups in the entire Gulf compared to our 3,422.

Because of discrepancies in method and loca-
tion of censusing, entire censuses are not very
useful for determining population change. How-
ever, comparisons of individual rookeries indi-
cate that the population in Mexico has increased.
For seven rookeries, our counts can be compared
with Lluch's (1969). Both the dates and method
of censusing were similar. He counted 5,977 ani-
mals and we counted 7,662 and 8,091 animals
in our two censuses. These represent increases
of 28 and 35% over a period of 13 to 16 years.
Our counts were higher on San Jorge (77%), San
Pedro Martir (56%), and Rocas Consag (181%),
and lower on Granito (40%), Roca Vela (81%),
and San Ignacio Farallon (alias Topolobampo)
(24%). Counts on San Pedro Nolasco were sim-
ilar in 1966 and 1979 but increased by 32% in
1981. Finally, it is not clear if the population
decreased from 1979 to 1981, because the two
censuses were conducted at slightly different times
during the breeding season. The 1 98 1 census was
made prior to peak season, accounting for the
lower count.

There are numerous difficulties in simply
counting large groups of sea lions, leading to
underestimates of animals present. Counts vary
with the census method used, weather condi-

CALIFORNIA SEA LION POPULATION IN MEXICO

TABLE 4. COMPARISON OF INDEPENDENT CENSUSES TAKEN AT SEA LION ROOKERIES IN THE GULF OF CALIFORNIA IN JUNE 1 98 1 .
Abbreviations: G = ground, L = launch, RC = R. Condit, FS = F. Sinsel, CF = C. Fox.

Rookery

Census
method

Weather, light,
terrain

Censuser

Males

Females

Pups

Juv.

Totals

Los Islotes

Clear sky, fading light;

rough terrain, boulder

& cobblestones.

San Pedro Martir

Windy, overcast; choppy

925

124

1,190

to heavy seas; 2 sandy

134

1,186

252

120

1,692

Los Machos (A.G.) L
(partial)

Granito (A.G.)

San Esteban

Totals

beaches, cobble coves,

rock tables, large boul-
ders backed by steep

cliffs.
Sunny, flat sea; most RC 84 832 226 247 1,389

females & pups on FS 67 1,069 293 109 1,535

sandy beaches, juve-
niles & subadult males

on rock tables & in

boulder fields,
s side calm; N side very RC 60 621 163 72 916

choppy, poor census- FS/CF 87 789 159 87 1,122

ing conditions; 1 sandy

beach on each side

bordered by irregular

rocky coastline backed

by steep cliffs.
Calm sea N & w sides; RC 234 1,865 369 500 2,968

steep cliffs with caves FS 212 1,824 545 160 2,741

& rocky shelves; long

sandy or cobbly

beaches.

C
FS/CF

458
516

4,269
4,882

837
1,255

943
482

6,507
7,132

tions, terrain, the experience and reliability of
censusers, and the size, age, and sex composition
of groups being counted. Moreover, the number
of sea lions on land varies with time of year and
the time of day. A special difficulty for estimating
breeding females is that some will always be at
sea feeding during the breeding season. As a result,
counts provide only minimum estimates of the
number of animals present. Empirically based
correction factors must be employed to estimate
actual population numbers.

We reason that the actual number of sea lions
counted in the Gulf of California underestimates
the number of animals and that a more valid
estimate is 20,144 animals. This estimate is
derived by applying three correction factors to
the 1979 total count of 15, 140 animals (the 1979
census is selected over the 1981 census because
it is closer to the peak of the breeding season).
This count is augmented by: (a) 744 animals,

representing known rookeries not censused in
1 979; San Ignacio Farall6n (323) and Rocas Con-
sag (421); (b) 3,496 pups, assuming that pups
counted reflect only 50% of the pups present; (c)
764 females, assuming that 10% of the females
are at sea feeding (see Bonnell et al. 1978). We
believe that 20, 1 44 total animals in the Gulf and
an annual production of approximately 7,000
pups are reasonable, conservative estimates.

Using the same assumptions, we can estimate
the number of sea lions on the Pacific coast of
Mexico. Before doing this, it should be noted
that previous censuses from launches or on foot
are of limited use for estimating population size
because they are only partial counts of the area
or they were conducted outside the breeding sea-
son, e.g., Bartholomew and Hubbs (1952), Rice,
Kenyon, and Lluch (1965), Orr, Schonewald, and
Kenyon (1970), Brownell, DeLong, and Schrei-
ber (1974). Aerial censuses of sea lions on the

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 7

TABLE 5. WEIGHTS AND MEASUREMENTS OF CALIFORNIA SEA LION PUPS OBTAINED 1 1-28 JUNE 1981. For each island listed,
the mean weight (in kilograms) or mean length (in centimeters) is shown plus or minus one standard deviation. N is in parentheses.

Weight

Length

Rookeries

Males

Females

Males

Females

San Pedro Martir

8.75 ± 1.89

7.50 ± 1.96

76.42 ± 4.58

71.43 ± 8.94

(6)

(4)

(6)

(4)

San Esteban

10.17 ± 1.54

8.00 ± 1.06

74.93 ± 3.00

74.08 ± 5.03

(6)

(9)

(6)

(9)

Granito (A.G.)

9.19 ± 1.04

7.75 ± 0.97

75.17 ± 2.62

71.48 ± 3.14

(16)

(13)

(16)

(13)

San Jorge

8.86 ± 0.92

7.71 ± 1.14

75.01 ± 2.30

72.55 ± 3.16

(9)

(12)

(9)

(12)

San Ignacio Farallon

8.80 ± 1.10

8.20 ± 1.30

76.20 ± 0.92

70.34 ± 3.31

(5)

Cedros

8.54 ± 1.45

7.29 ± 1.93

76.79 ± 3.37

72.82 ± 3.91

(12)

(14)

(12)

(14)

Totals

9.01 ± 1.32

7.60 ± 1.45

75.71 ± 2.93

72.34 ± 4.18

(54)

(57)

(54)

(57)

Pacific coast of Mexico by Mate (1977) (see Bon-
nell et al. 1978) yielded a direct count of 45,872
animals. We estimate that the total number of
sea lions on the Pacific coast of Mexico is approx-
imately 63,020 animals, a number which includes
the annual production of 29,000 pups. We start
with the direct count of 23,098 animals obtained
in 1979. We increase this number by: (a) 4,777
representing two rookeries we counted in 1981
but not in 1979 (Islas Cedros and Magdalena,
see Table 2); (b) 12,682 representing the follow-
ing rookeries we did not census but which were
censused by Mate (1977) (see also Bonnell et al.,
1978): Los Coronados (297), San Geronimo
(1,113), Punta Rosarito (2,722), Natividad
(5,785), and Punta Tosca (2,765); (c) 8,667 pups,
assuming that 50% of the pups counted from
launches were missed; (d) 1 1,445 pups, to com-
pensate for the failure to count pups from aerial
censuses; and (e) 2,351 females, assuming that
1 0% of the females were at sea. Augmentations
in categories b, d, and e are based on Mate's aerial
census in 1975 (Mate 1977). Since he did not
categorize animals by age or sex, we added pup
and female counts to his numbers using ratios
derived from our censuses.

From these estimates and censuses conducted
in the United States, we can estimate the size of
the population. Bonnell et al. (1978) estimated
50,000 animals in southern California waters
from aerial censuses and 4,000 in the northern
part of the nonbreeding range. Thus, the total

number of California sea lions in the United States
and Mexico is approximately 145,000 animals.
Of this total, 16% are in the Gulf of California,
46% are on the Pacific coast of Mexico, 35% are
in southern California, and 3% range as far north
as Vancouver Island, British Columbia (Han-
cock 1970; Bigg 1973). Our estimate of the United
States and Mexico population of this species
exceeds the figure of 125,000 estimated by Bon-
nell et al. (1978). Both estimates do not include
the geographically separated subspecific popu-
lations on the Galapagos Islands and in Japan.
The distribution of breeding animals in the
Gulf differs from that most commonly observed
on Pacific coast rookeries. In the Gulf, male ter-
ritories are predominantly aquatic, fronting on
small groups of females and pups distributed in
a narrow zone along the water's edge on cobble-
stone beaches, among boulders, or on rocky
ledges. On Pacific coast rookeries in Mexico and
California, females are most often hauled out in
large groups well above the surf line on sandy
beaches or on flat rock outcroppings sloping into
the sea (Peterson and Bartholomew 1967; Odell
1975; Bonnell et al. 1978); male territories are
semi-aquatic or usually terrestrial. Higher tem-
peratures in the Gulf may account in part for
these differences in behavior and distribution.

ACKNOWLEDGMENTS

We thank Drs. Felix Cordoba Alva and Carlos
de Alba Perez for help in obtaining permits and

CALIFORNIA SEA LION POPULATION IN MEXICO

making logistical arrangements in Mexico; George
Shor, Robert Haines, and others at Scripps Insti-
tution of Oceanography for helping to make the
R/V Ellen B. Scripps available to us; Captains
Hansen and Whitman and their crews for safe
and efficient conduct; and Dr. Leo Ortiz, Dr.
James Estes, Dr. Martha Field, Dr. Robert Brow-
nell, Jr., Steve Davenport, Edward Keith, Kathy
Panken, John Peterson, Joanne Reiter, Marianne
Riedman, and Keith Skaug for field assistance.
These expeditions were supported in part by
National Science Foundation grant DEB 77-
17063 AO1 to B. Le Boeuf.

LITERATURE CITED

AURICLES, D., C. Fox, AND F. SINSEL. 1981. Species of fishes
identified with the otoliths found in scats of sea lions at "Los
Islotes" Island, B.C.S., Gulf of California, Mexico. Abstract,
4th Biennial Conference on the Biology of Marine Mammals,
Dec. 14-18, 1981, San Francisco, California.

, R. ROMERO, AND C. Fox. 1978. Censos poblaciones

de lobos marines (Zalophus californianus) por edades y sexos
en la bahia de la Paz y sus alrededores. Pages 139-154 in
Informe General de Labores, 1978. Centre de Investiga-
ciones Biologicas de Baja California, A.C.

BARTHOLOMEW, G. A., AND C. L. HUBBS. 1952. Winter pop-
ulations of pinnipeds about Guadalupe, San Benito, and
Cedros Islands, Baja California. J. Mammal. 33:160-171.

BIGG, M. A. 1973. Census of California sea lions on southern
Vancouver Island, British Columbia. J. Mammal. 54:285-
287.

BONNELL, M. L., B. J. LE BOEUF, M. O. PIERSON, D. H. DETTMAN,

AND G. D. FARRENS. 1978. Pinnipeds of the southern Cal-
ifornia Bight. Vol. Ill, Part 1, 535 p. in K. S. Norris, B. J.
Le Boeuf, and G. L. Hunt, Jr., eds., Marine Mammal and
Seabird Surveys of the Southern California Bight Area, 1975-
1978. Bureau of Land Management, Dept. of Interior Con-
tract AA550-CT7-37, Government Printing Office, Wash.,
D.C.

BROWNELL, R. L., JR., R. L. DE LONG, AND R. W. SCHREIBER.
1974. Pinniped populations at Islas de Guadalupe, San Be-
nito, Cedros and Natividad, Baja California, in 1968. J.
Mammal. 55:469^172.

HANCOCK, D. 1970. California sea lion as a regular visitant
off the British Columbia coast. J. Mammal. 51:614.

LLUCH, B. D. 1969. El lobo marino de California (Zalophus
californianus californianus Lesson, 1828) (Allen, 1880), en
dos mamiferos marines de Baja California. Inst. Mex. Rec.
Nat. Renovables., 3-69.

MATE, B. 1977. Aerial censusing of pinnipeds in the Eastern
Pacific for assessment of population numbers, migratory dis-
tributions, rookery stability, breeding effort, and recruit-
ment. Report to Marine Mammal Commission Contract
MM5AC001, 28 Feb. 1977. 68 p.

ODELL, D. K. 1975. Breeding biology of the California sea
lion, Zalophus californianus. Rapp. P.-V. Reun. Cons. Int.
Explor. Mer 169:374-378.

ORR, R. T., J. SCHONEWALD, AND K. W. KENYON. 1970. The
California sea lion: skull growth and a comparison of two
populations. Proc. Calif. Acad. Sci., ser. 4, 37:381-394.

PETERSON, R. S., AND G. A. BARTHOLOMEW. 1967. The nat-
ural history and behavior of the California sea lion. Spec.
Pub. No. 1, Amer. Soc. Mammal. 79 p.

RICE, D. W., K. W. KENYON, AND D. LLUCH. 1965. Pinniped
populations at Islas Guadalupe, San Benito, and Cedros,
Baja California, in 1965. Trans. San Diego Soc. Nat. Hist.
14:73-84.

CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, California 94 1 1 8

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 8, pp. 87-110, 15 figs. July 6, 1983

MEGAMOUTH-A NEW SPECIES, GENUS, AND FAMILY OF

LAMNOID SHARK (MEGACHASMA PELAGIOS, FAMILY

MEGACHASMIDAE) FROM THE HAWAIIAN ISLANDS

Leighton R. Taylor

Waikiki Aquarium, 1777 Kalakaua Ave., Honolulu, Hawaii 96815

L. J. V. Compagno

Tiburon Center for Environmental Studies, San Francisco State University,
P.O. Box 855, Tiburon, California 94920

and
Paul J. Struhsaker

1050 Koloa St., Honolulu, Hawaii 96816

ABSTRACT: Megachasma pelagios, a new genus and species of lamnoid shark assigned to the new family
Megachasmidae, is described and defined from a single adult male, 4.46 m total length. The holotype and
only known specimen was collected approximately 42 km NE of Oahu, Hawaii. Structure and habitus distinctly
differ from other lamnoid sharks, particularly in head and tooth morphology and in mesopelagic filter feeding.

INTRODUCTION 4) Crew members of the AFB-14 realized that

On 15 November 1976, the research vessel the shark was unusual and brought it aboard with

AFB- 14 of the Naval Undersea Center (now the much difficulty. The shark was shipped to the

Naval Ocean Systems Center), Kaneohe, Hawaii, Kaneohe Bay facility of the Naval Undersea Cen-

was conducting oceanographic research in waters ter and tied alongside the dock overnight,
about 42 km northeast of Kahuku Point, Oahu, The senior author inspected it the next morn-

at about 21°51'N and 157°46'W. From 1015 to ing. Preliminary examination indicated that it

1415 Hawaiian Standard Time the ship had de- represented a very distinct, undescribed species,

ployed two large parachutes as sea anchors at a and it was decided that it should be preserved

depth of about 165 m in water with a bottom intact. Accordingly, the shark was winched out

depth of approximately 4600 m. When the para- of the water by the tail using a Navy crane, but

chutes were hauled to the surface, using a small the caudal fin broke off and the shark fell into

winch with an 1 80 kg pull shut-off, one of them the water and had to be retrieved by divers. The

had entangled in it a large adult male shark 4.46 shark was quick-frozen at Hawaiian Tuna Pack-

m (14.6 ft) long and 750 kg (1653 Ibs) (Figs. 1- ers, Honolulu, while a large preservation tank

[87]

FIGURE 1 . Artist's conception of Megachasma pelagios in its natural habitat, slowly swimming with open mouth and feeding
on planktonic animals. (From a painting by Richard Ellis.)

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

was constructed. On 29 November 1976, the
shark was transported frozen to the Kewalo dock
site of the National Marine Fisheries Service for
thawing and injection with formalin. Subsequent
examination of the shark by the authors and col-
leagues indicated that it is a lamniform shark
(order Lamniformes of Compagno 1973a) that
is not assignable to any known genus or family
and is herein described as Megachasma pelagios,
new genus and species, and placed in the new
family Megachasmidae.

The discovery of the novel shark was widely
reported in newspapers (e.g., Anonymous 1 and
2, 1976, and Dunford 1976). It was dubbed the
"Megamouth shark" in reference to its unusually
large oral cavity. This common name has since
been adopted by several authors (Compagno
1977, 1979, and 1981; Taylor 1977; Tinker 1978;
Faughnan 1 980; Clark 1981), and we suggest that
it be considered as the accepted common name
for the species.

This strange shark is extraordinary in its dis-
tinctness from other sharks and its great size.
Most sharks are small, less than 2 m long at
maturity (Compagno 198 1). The new shark joins
the company of the few giant sharks commonly
reaching total lengths over 4 m, including the
broadnose sixgill shark (Hexanchus griseus), Pa-
cific sleeper shark (Somniosus pacificus), Green-
land shark (S. microcephalus), whale shark (Rhi-
niodon typus), great white shark (Carcharodon
carcharias), tiger shark (Galeocerdo cuvier), and
great hammerhead (Sphyrna mokarrari). The
common thresher (Alopias vulpinus) and bigeye
thresher (A. superciliosus) also reach total lengths
over 4 m, but these sharks have greatly elongated
caudal fins and hence are relatively small-bodied
in comparison to the giant species. Although new
species of small sharks are discovered fairly fre-
quently, giant sharks are not, and almost all of
the great species were described in the 1 8th and
19th centuries.

Because the only known specimen of Mega-
chasma pelagios is an adult male, and because
it is very common for female sharks to reach a
somewhat larger size than males, it is reasonable
to expect larger specimens of this species.

METHODS

On 30 November 1976, the thawed shark was
placed in a large, above-ground plastic pool filled
with seawater. Comprehensive measurements of
the shark were recorded, following the proce-

dures of Bigelow and Schroeder (1948). Skin
samples were taken from the mouth, tongue, pec-
toral fin, caudal fin, back below first dorsal fin,
and gill-rakers for later examination using a
Cambridge S410 Stereoscan electron micro-
scope.

Skin samples from the mouth lining and tongue
were sectioned and stained using standard his-
tological techniques. A short incision, approxi-
mately 30 cm long, was made on the ventral
surface to gain access to the stomach and val-
vular intestine, and stomach contents were re-
moved. The valvular intestine was removed, slit
medially to count the ring valves and to remove
intestinal worms for parasitologists (Dailey and
Vogelbein 1 982), and separately preserved. Sam-
ples of muscle tissue and liver were taken for
electrophoretic analysis. Extensive sets of still
photos were made of the preservation process by
the authors and Mr. Paul Meyers of the Naval
Undersea Center, who also made 1 6 mm movies
of these techniques.

The body cavity and musculature of the shark
were injected with 25 1 of 100% formalin (40%
aqueous formaldehyde gas solution). The shark
was then lifted by crane and cargo sling into a
4 X 3.5 X 1 m fiberglass box and covered with a
40% sea water-formalin solution. After six months
in formalin, the specimen was deposited in the
fish collection of the Bernice P. Bishop Museum,
Honolulu, where it was rinsed for 30 days in
water and then placed in 55% isopropyl alcohol.
Tooth samples were removed for examination
of their morphology, and one tooth was sectioned
and stained for tooth histology. A "peel" dissec-
tion was made on the right side of the head to
examine the neurocranium and jaw structure of
the shark, and similar dissections were made on
the right pectoral fin and right clasper. Vertebrae
were excised from the base of the caudal fin and
from beneath the first dorsal fin and sectioned
to examine their calcification patterns.

Terminology for descriptive morphology of
Megachasma pelagios follows Bigelow and
Schroeder (1948) and Compagno (1970, 1973a,
1973b, and 1979).

Megachasmidae, new family

TYPE-GENUS.— Megachasma Taylor, Compagno, and Struh-
saker, new genus.

FAMILY DESCRIPTION. — Giant neoselachian
sharks of the order Lamniformes (as defined by

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8

FIGURE 2. Holotype of Megachasma pelagios, within 12 hours of its capture. Note the extreme protrusibility of the jaws
and the gill filaments visible in the first gill opening.

Compagno 1973a) reaching at least 4.46 m length
when adult. Trunk cylindrical but not highly fu-
siform, tapering rearward from the head. Caudal
peduncle short, stout, slightly compressed, and

without lateral keels or ridges; a shallow, longi-
tudinally oval upper precaudal pit present, but
no lower pit. Head broad, very large and long,
and not pointed, length greater than abdomen

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

FIGURE 3. Frontal view. Note Navy research vessel and winch which retrieved shark in background. (Official U.S. Navy
photograph.)

between pectoral and pelvic bases. Snout very
short, depressed, and broadly rounded, not con-
ical or bladelike. Eyes lateral on head, length less
than one-fourth length of longest gill openings.

Nostrils small, widths about '/,, internarial
width, with short, low anterior nasal flaps; nos-
trils lateral and opposite the first fourth of mouth.
Gill openings moderately large, not expanded

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8

onto dorsal surface of head; internal gill openings
with numerous gill-rakers of a unique type,
formed as elongated, slender, cartilage-cored
dermal papillae covered by imbricated denticles
(Fig. 6). Mouth terminal and very large, broadly
arched, extremely long, and extending far behind
eyes when jaws are not protruded; jaws strongly
protrusible, capable of extension well in front of
snout. No true labial furrows or labial cartilages,
but with inner labial grooves present along edges
of mouth corners.

Teeth similar in upper and lower jaws, weakly
differentiated, with moderately long, broad, flat-
tened roots, very short labial root lobes, and very
long, broad, expanded lingual protuberance;
moderately strong basal ledges and grooves on
the labial crown face; a broad, enameloid-free
neck on the crown foot; a strong, narrow, lin-
gually hooked cusp with cutting edges confined
to its tip, no striations or ridges on the cusp.
Teeth not compressed and bladelike, relatively
small, and very numerous, over 100 rows in each
jaw and in three or four functional series. Tooth-
less spaces on symphyses of jaws extremely broad,
especially on upper jaw. Teeth in each jaw half
apparently continuously varying and without
discrete row groups; no gap or reduced inter-
mediate teeth between teeth in anterior and lat-
eral positions in upper jaw.

Lateral trunk denticles with broad, teardrop,
or wedge-shaped, flattened crowns, not erect,
hooked, or directed anteriorly or dorsoventrally;
pedicels of trunk denticles low and broad (Fig.
1 1). Wavy grooves of naked skin on the pectoral,
pelvic, and caudal fin webs. Pectoral fins rela-
tively narrow, long and blunt-tipped, length from
origin to free rear tip about half as long as pec-
toral anterior margin. Origins of pectoral fins un-
der fourth gill openings. Pectoral fins more than
three times area of first dorsal fin, with anterior
margins more than three times length of pelvic
anterior margins. Pectoral fin skeleton plesodic,
with pectoral radials extending into the distal fin
web nearly to its edge; ceratotrichia reduced along
distal fin margin and not extending proximally
to radial musculature of fin. Pectoral fins very
small, angular, smaller than first dorsal fin but
larger than second dorsal, with an aplesodic fin
skeleton. Claspers moderately slender and elon-
gated, with attenuated tips and external spurs
(Fig. 1 1 ). First dorsal fin moderately large, angular
and relatively low, with a narrowly rounded apex

and an aplesodic fin skeleton; origin of first dorsal
much closer to pectoral fin bases than pelvic bas-
es, and free near tip, well in front of pelvic origins.
Second dorsal fin less than one-third area of first
dorsal and slightly less than half as high, angular
and broad-based, with its origin about over the
pelvic fin insertions. Neither second dorsal nor
anal bases pivoted. Anal fin about half area of
second dorsal, angular and broad-based, with its
origin about opposite free rear tip of second dor-
sal and its free rear tip well in front of ventral
caudal origin; insertion of anal separated from
ventral caudal origin by space greater than base
or anal. Caudal fin with a long dorsal lobe nearly
half length of rest of shark, a long ventral lobe
about % as long as dorsal lobe, a deeply notched
postventral caudal margin, a weak subterminal
notch, and no undulations or ripples on the dor-
sal or preventral caudal margins; caudal fin not
lunate or crescentic, dorsal caudal vertebral axis
moderately elevated at an angle to body axis (het-
erocercal).

Neurocranium (Fig. 1 3) with tripodal rostrum
formed of a small, moderately elongated, medial
rostral cartilage originating from expanded in-
ternasal plate and pair of basally enlarged, tri-
angular lateral rostral cartilages that taper an-
teromedially to fuse with medial rostral cartilage
and form a narrow, flattened, unfenestrated ros-
tral node. Base of medial rostral cartilage ele-
vated by dorsally arched internasal septum above
level of bases of lateral rostral cartilages and nasal
capsules, so that medial rostral cartilage arches
anteroventrally to meet rostral node. Rostrum
short, less than half nasobasal length of cranium.
Nasal capsules small, greatly compressed, far lat-
eral to each other and separated by flattened in-
ternasal septum. Entire anterior surfaces of nasal
capsules forming bases of lateral rostral carti-
lages. Broad subethmoid fossa not extending an-
terior to nasal capsules. Cranial roof very broad
and flat, not arched above the orbits, with a huge
transverse anterior fontanelle; basal plate broad-
ly arched. Orbits with low preorbital processes,
complete supraorbital crests, and broad, low
postorbital processes. A deep pit on each side of
ventral surface of cranium between base of sub-
orbital shelf and basal plate in front of stapedial
fenestrae, for orbital processes of palatoquad-
rates. Otic capsules broad and relatively long,
without elongated pterotic horns.

Jaws very long and stout, much longer than

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

cranium, extending from the cranial rostral node
to well behind the occiput when retracted. Pal-
atoquadrates with long, stout palatine processes
lacking dental bullae; strong, low, knoblike, car-
tilaginous orbital processes, and low, strong
quadrate processes. Orbital processes articulat-
ing on ventral surfaces of suborbital shelves and
basal plate below orbits, quadrate processes far
below postorbital processes of cranium and not
contacting them. Anterior ends of Meckel's car-
tilages extending below level of anterior ends of
palatoquadrates, no "overbite" of palatoquad-
rates on Meckel's cartilages. Vertebral column
with well-developed cartilaginous centra sepa-
rated by broad bands of notochordal sheath, but
with primary and secondary calcification vir-
tually absent. Intestinal valve with 24 turns.

CLASSIFICATION.— Compagno (1973a, 1977)
divided the living elasmobranch fishes, or neo-
selachians, into four superorders, of which the
Galeomorphii or galeomorph sharks clearly in-
cludes the new family Megachasmidae and genus
Megachasma. Megachasmidae has the following
galeomorph characters: head and body not great-
ly depressed and not expanded laterally; spiracles
without valves; five pairs of laterally situated gill
openings; denticles covering almost entire body,
not absent ventrally, nor enlarged on midline of
back, and not enlarged on pectoral fins in adult
males; pectoral fins without anteriorly expanded
triangular lobes covering gills or fused to sides
of head above them; propterygium of pectoral
fin skeleton not anteriorly expanded; pectoral fins
not modified into propulsive organs; pectoral gir-
dle not articulating with vertebral column; vent
confluent with pelvic fins; two dorsal fins and an
anal fin present; caudal fin heterocercal, with a
subterminal notch on the dorsal caudal lobe and
with ventral lobe shorter than dorsal lobe; neuro-
cranium with strong suborbital shelves, no antor-
bital cartilages, ectethmoid processes, or en-
larged ectethmoid chambers on the nasal capsules,
rostrum not trough-shaped, no basal angle on
basal plate, no lateral commissures on otic cap-
sules, and with incomplete postorbital walls; no
palatobasal articulation of palatoquadrates with
neurocranium; hyoid arch complete, no pseu-
dohyoids; vertebral column without synarcuals,
and vertebral centra without concentric calcifi-
cations.

The Galeomorphii of Compagno ( 1 973a, 1 977)

was subdivided into four orders, Heterodonti-
formes, Orectolobiformes, Carcharhiniformes,
and Lamniformes; of these, the family Mega-
chasmidae falls in the order Lamniformes or
lamnoid sharks. Lamnoid characters of Mega-
chasmidae include its simple nostrils of the or-
dinary shark type, entirely separate from the
mouth, with small anterior nasal flaps, diagonal
incurrent and excurrent apertures, and no peri-
nasal folds and grooves, anterior barbels, or na-
soral grooves; a long mouth extending behind
the eyes when jaws are retracted; no supraorbital
and subocular ridges; eyes circular and laterally
without nictitating eyelids, subocular pouches,
or postorbital eyelid muscles; osteodont teeth (Fig.
9) with weak basal ledges; posterior teeth not
enlarged and formed into molariform crushers;
claspers with external spurs on the T-3 cartilage
and with elongated, tubular, expanded marginal
cartilages; dorsal fins spineless, with segmented
basal cartilages; cranium with a tripodal rostrum,
nasal capsules not anteroposteriorly elongated and
trumpet-shaped; no ethmopalatine grooves for
the articulation of the palatoquadrate orbital
processes, complete preorbital walls, separate fo-
ramina for superficial ophthalmic nerves in or-
bits and for hyomandibular nerves on otic cap-
sules, and relatively long otic capsules; jaws long,
extending posterior to the occiput; mouth gape
not restricted anteriorly, labial cartilages, folds
and grooves reduced or absent; pectoral fin skel-
eton with a small propterygium, moderately large
mesopterygium, and large metapterygium; me-
sopterygium and metapterygium not elongated
parallel to the axes of their radials, and not proxi-
mally shaftlike, distally expanded and without a
fenestra between them; preorbitalis or levator
labii superioris muscles relatively small and
anteroposteriorly positioned on the jaws, with
origins on posteroventral surfaces of the nasal
capsules, fibers nearly horizontal when jaws are
retracted, and insertions far posterior on the ad-
ductor mandibulae muscles at the jaw angles;
adductor mandibulae muscles anteriorly notched;
levator palatoquadrati muscles simple, not sub-
divided into anterior constrictor and spiracular
muscles; no craniomandibular or mandibulo-
cutaneous muscles; and an elongated, ring- valve
intestine.

RELATIONSHIPS TO OTHER LAMNOIDS.— Within
the Lamniformes, the family Megachasmidae

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8

represents a very distinct and singular taxon,
well separated from all other families. Other
lamnoids of the families Odontaspididae, Mitsu-
kurinidae, Pseudocarchariidae, Alopiidae, Ceto-
rhinidae, and Lamnidae all differ from the Me-
gachasmidae in having a more elongated, more
narrowly rounded, conical or bladelike snout; no
papillose gill-rakers (denticle gill-rakers present
in Cetorhinidae); mouth subterminal on head and
less enlarged; tongue smaller; upper anterior and
lateral teeth separated by a gap that may or may
not have reduced intermediate teeth; tooth rows
either less than 60 in each jaw, or more than 200
(Cetorhinidae); toothless space on upper sym-
physis relatively narrow; no wavy grooves of na-
ked skin on the pectoral, pelvic, and caudal fins;
lateral rostral cartilages narrow-based and only
covering part of the dorsal surfaces of the nasal
capsules or the preorbital processes; base of me-
dial rostral cartilage well below bases of lateral
rostral cartilages and with shaft of cartilage below
rostral node; nasal capsules nearly spherical, not
compressed, and with ventral nasal apertures;
cranial roof narrow to only moderately expanded
anteriorly, with anterior fontanelle varying from
moderate to greatly reduced; orbital processes
more or less reduced on palatoquadrates, artic-
ulating with the suborbital shelves where present;
jaws shorter, beginning well behind the snout tip
when retracted; and with primary calcification
of the double cones and secondary radii well de-
veloped in their vertebral centra.

Members of the family Odontaspididae (in-
cluding the genera Eugomphodus and Odontas-
pis) further differ from Megachasma and the Me-
gachasmidae in having prominent, transverse
precaudal pits; labial folds, furrows and carti-
lages present (with the possible exception of E.
tricuspidatus); nostrils in front of the mouth; teeth
with strong labial root lobes, moderate lingual
protuberances, narrow necks on the crown, and
labiolingually diagonal attachment surfaces;
symphyseal, anterior, lateral, intermediate, and
posterior tooth-row groups well differentiated
along dental bands, with anteriors and laterals
enlarged; pectoral fins smaller, shorter, broader,
less elongated, and not falcate, and with aple-
sodic fin skeletons; pectoral fin origins behind
fifth gill openings; claspers stouter and blunt-
tipped, with blunt clasper spurs; origin of first
dorsal fin well posterior to pectoral insertions;

second dorsal fin more than half as high as first
dorsal; caudal fin shorter, less than half as long
as rest of shark; subterminal notch of caudal fin
deep; ventral caudal lobe shorter, dorsal caudal
margin with rippled edges; rostral node com-
pressed, with vertical fenestra and strut; cranial
roof narrow and arched above orbits; and otic
capsules with strong pterotic horns.

The family Pseudocarchariidae, which like
Megachasmidae has a single, oceanic, highly dis-
tinct species (Pseudocarcharias kamoharai), dif-
fers from Megachasma in many characters, in-
cluding its more slender body and shorter head;
slender, cylindrical caudal peduncle with low lat-
eral keels and upper and lower transverse, cres-
centic precaudal pits; much larger eyes; nostrils
anterior to mouth; more elongated gill openings,
extending onto dorsal surface of head; teeth with
strong labial root lobes, moderate lingual pro-
tuberances, a narrow neck on the crown, and
labiolingually diagonal attachment surfaces; an-
teriors, intermediates, and lateroposteriors well
differentiated in dental bands; anteriors and an-
terior-laterals enlarged, pectoral fins smaller,
broader, less elongated, and not falcate, with
aplesodic fin skeletons; origins of pectoral fins
behind fifth gill openings; anal fin with a narrow
base and pivotable; caudal fin with a shorter dor-
sal and ventral caudal lobe; rostrum longer, with
appendices, a compressed rostral node, and ver-
tical fenestrae and struts; basal plate and cranial
roof extremely narrow, with narrow, slotlike,
vertical anterior fontanelle; orbits of cranium very
large; pterotic horns present and well developed
on otic capsules; palatine processes of palato-
quadrates enlarged and forming large dental bul-
lae, articulating with the orbital notches of the
cranium; and quadrate processes of palatoquad-
rates elevated and contacting postorbital pro-
cesses.

The benthopelagic family Mitsukurinidae also
has a single living, strongly distinct species (Mit-
sukurina owstoni). The Mitsukurinidae differs
from the Megachasmidae in lacking precaudal
pits; having a greatly elongated, flattened, blade-
like snout; smaller eyes; a very narrow, elongated
mouth; lower labial furrows; teeth with strong
labial root lobes, moderate lingual protuber-
ances, a narrow neck and striations on the crown,
and labiolingually diagonal attachment surfaces;
symphyseals, anteriors, laterals, and posteriors
well differentiated in dental bands, with anteriors

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

and laterals enlarged; lateral trunk denticles with
narrow, hooked, semierect crowns; pectoral fins
smaller than pelvic fins, shorter, broader, not
elongated and falcate, and with aplesodic fin skel-
etons; pectoral origins behind fifth gill openings;
first and second dorsal fins equal-sized, smaller
than pelvic and anal fins; anal fin large, broadly
rounded, and separated from lower caudal origin
by a narrow notch; anal fin origin about opposite
or close behind second dorsal origin; no ventral
caudal lobe; rostrum of cranium greatly elon-
gated, longer than nasobasal length of cranium,
with a compressed, extremely long rostral node;
subethmoid fossa extending anterior to the nasal
capsules; supraorbital crest reduced to separate
preorbital and postorbital processes; and with
palatine processes of palatoquadrates deflected
ventrally, with prominent bullae.

The three highly specialized lamnoid families
Alopiidae, Lamnidae, and Cetorhinidae have
numerous additional differences from the Me-
gachasmidae. The Alopiidae further differs from
the Megachasmidae in having a shorter head;
crescentic upper precaudal pits; larger eyes; nos-
trils anterior to mouth; shorter gill openings; a
much smaller mouth and less highly protrusible
jaws; teeth with weaker lingual protuberances,
stronger labial root lobes, and differentiated an-
teriors, lateroposteriors, and (variably) inter-
mediates and symphyseals; claspers very slender,
without spurs; pelvic fins plesodic; first dorsal fin
higher and plesodic, with its origin well posterior
to the pectoral insertions; second dorsal much
smaller relative to first dorsal, with a narrow,
pivotable base; anal fin smaller, with narrow,
pivotable base; caudal fin about as long as rest
of shark, with a rippled dorsal margin; rostral
node of rostrum compressed, with a vertical fe-
nestra and strut; internasal septum narrow and
high; subethmoid fossa very narrow; cranial roof
narrow, flat or strongly arched; orbits large to
gigantic; and palatine processes of palatoquad-
rates with small dental bullae.

The Lamnidae differs from the Megachasmi-
dae in the following additional characters: trunk
more fusiform; caudal peduncle greatly de-
pressed, with strong lateral keels, and with trans-
verse, crescentic, upper and lower precaudal pits;
nostrils anterior to the mouth; gill openings long-
er, extending partway onto dorsal surface of head;
jaws less protrusible; teeth with low lingual pro-
tuberances, enlarged anteriors, laterals, and in-

termediates; pectoral fin origins behind fifth gill
openings; second dorsal much smaller relative to
first dorsal, with a narrow, pivotable base; anal
fin slightly larger than second dorsal, with a nar-
row, pivotable base; caudal fin shorter, less than
half length of rest of shark, nearly symmetrical
and lunate in Lamnidae, with a relatively shorter
dorsal lobe, ripples in dorsal margin, and a longer
ventral lobe; cranial roof narrow and arched; otic
capsules with elongated pterotic horns; palatine
processes of palatoquadrates with prominent
dental bullae articulating with underside of eth-
moid region of cranium; and quadrate processes
of palatoquadrates very high.

Finally, the family Cetorhinidae with the only
other filter- feeding lamnoids of the genus Cetor-
hinus, differs from the family Megachasmidae in
the following particulars: trunk more fusiform;
caudal peduncle somewhat depressed, with strong
lateral keels and transverse, crescentic upper and
lower precaudal pits; nostrils anterior to mouth;
gill openings much larger, expanded onto dorsal
and ventral surfaces of head; jaws little protru-
sible; pectoral fins with their origins behind fifth
gill openings; lateral trunk denticles with erect,
hooked, narrow crowns, directed anteriorly and
dorsoventrally as well as posteriorly; claspers
stout, with broad tips and heavy spurs; first dor-
sal fin with its origin far posterior to pectoral
insertions, and midbase closer to pelvic bases
than to pectoral bases; caudal fin shorter, less
than half length of rest of shark, nearly sym-
metrical and lunate, with a shorter dorsal lobe
and longer ventral one; medial ros'tral cartilage
very broad, platelike, and ventrally excavated by
the broad anterior expansion of the subethmoid
fossa; lateral rostral cartilages joining each other
posterior to their junction with the rostral node,
and extending anterior to that junction as a me-
dial rod; cranial roof moderately broad, highly
arched above orbits; supraorbital crests fenes-
trate basally; and jaws very slender and weak.

The phenetic comparisons between Mega-
chasmidae and other lamnoids presented above
are not intended to be exhaustive, but serve to
demonstrate the separation of Megachasmidae
from related families. They do not broach the
question of the relationship of the megamouth
shark to other lamnoids. A detailed account of
lamnoid interrelationships is beyond the scope
of this paper, but suffice it to note here that many
of the characters of Megachasma pelagios, such

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8

as its snout and jaw structure, gill-rakers, dermal
grooves on fins, reduced vertebrae, and ethmoid
morphology are evidently unique derived char-
acters of this shark that do not offer a clue to its
relationships. Its teeth are superficially similar
to those of the basking shark (Cetorhinus max-
imus), but this may be parallel evolution of ves-
tigial structures in two very different lamnoid
filter-feeders (as suggested by the superficial sim-
ilarity of the teeth of the orectoloboid whale shark,
Rhiniodon typus). Megachasma most resembles
members of the Odontaspididae (especially the
genus Odontaspis) and Pseudocarchariidae in its
body shape, fin shape, relative fin sizes (except
for the pectoral fins), fin positions, and relation-
ships of interspaces between fins to fin size. By
comparison with the derived families Alopiidae,
Cetorhinidae, and Lamnidae, these similarities
between Megachasmidae, Odontaspididae, and
Pseudocarchariidae may prove to be common
primitive characters not of importance in dem-
onstrating phyletic relationships among these
families. Megachasma shares the derived char-
acter state of plesodic pectoral fins with the Al-
opiidae, Cetorhinidae, and Lamnidae, but pres-
ently appears to have little else in common with
these derived families.

On the other hand, two characters of Mega-
chasma, if correctly interpreted as primitive, sug-
gest that Megachasmidae is the sister-group of
all other living lamnoids. The absence of differ-
entiated anteriors, laterals, and intermediates (or
a toothless gap between anteriors and laterals) in
Megachasma may indicate that it is primitive in
lacking them, and that all other lamnoids (in-
cluding Cetorhinus) can be united by the pres-
ence of these tooth-row groups as a shared de-
rived character. However, the unusually broad,
toothless space at the upper symphysis of Me-
gachasma suggests another possibility, that it is
derived in having lost these row groups, at least
in the upper jaw; and that the simple gradient
monognathic heterodonty in the dentral bands
is secondary and correlated with the evolution
of gill-rakers as the primary feeding structures in
Megachasma.

The second character is the well-developed or-
bital processes on the palatoquadrates of Me-
gachasma, which suggest a primitive condition
by comparison with other, non-lamnoid sharks.
The reduced (Alopiidae, Odontaspididae, Mit-
sukurinidae, and Cetorhinidae) or apparently

nonexistent (Lamnidae, Pseudocarchariidae) or-
bital processes of other lamnoids would by this
interpretation represent a shared derived char-
acter of lamnoids other than Megachasma. A
detailed assessment of these characters and oth-
ers, grouping the various lamnoid genera and
families will be considered in detail elsewhere
(Compagno, in preparation).

A possible fossil relative of M. pelagios is rep-
resented by isolated small teeth (2-15 mm high)
known since the 1960's from early Miocene de-
posits in the southeastern San Joaquin Valley of
California (Shelton P. Applegate, pers. comm.),
and subsequently found in other localities in the
late Oligocene or early Miocene of northern Cal-
ifornia (Phillips et al. 1976) and central Oregon
(Bruce J. Welton, pers. comm.). The shark rep-
resented by these teeth has never been named,
but is known from abundant tooth material from
southern California. Its affinities have been much
debated among palaeoichthyologists, but it ap-
pears most likely to be a lamnoid because of its
osteodont tooth histology and external tooth
morphology. Dr. Bruce J. Welton is preparing a
paper describing this shark, and will compare it
with M. pelagios, of which it is possibly a fossil
congener but is distinctly more primitive.

Megachasma, new genus

TYPE-SPECIES.— Megachasma pelagios Taylor, Compagno, and
Struhsaker, new species.

DERIVATION OF NAME. — mega, from Greek,
large, great; chasma, yawning hole, open mouth.

GENERIC DIAGNOSIS.— Characters of the new
genus are those of the new family Megachas-
midae (see above).

Megachasma pelagios, sp.nov.

MEGAMOUTH SHARK

HOLOTYPE.— An adult male, 4460 mm total length, Bernice
P. Bishop Museum, Honolulu, Oahu, Hawaii, BPBM 22730.

TYPE-LOCALITY.— Hawaiian Islands, about 42 km NE Ka-
huku Point, Oahu, 21°51'N, 157°46'W, at about 165 m depth
in water about 4600 m deep.

DERIVATION OF SPECIES NAME.— pelagios, from
Greek, of the open sea.

MEASUREMENTS AND PROPORTIONS.— These are
given below as measurements in millimeters, fol-
lowed by their proportions as percentages of total
length and precaudal length, given in that order
in parentheses.

Total length: 4460 mm (100% total length,
144.3% precaudal length).

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

FIGURE 4. Lateral-view drawing of the holotype of Megachasma pelagios, with jaws in retracted position. Drawn by L. J.
V. Compagno.

Precaudal length (snout to upper caudal ori-
gin): 3090(69.3, 100).

Tip of snout to: upper symphysis, 66 (1.5, 2.1);
nostrils, 100 (2.2, 3.2); orbits, 240 (5.4, 7.8); spi-
racles, 450 (10.1, 14.6); 1st gill openings, 850
(19.1, 27.5); 2nd gill openings, 920 (20.6, 29.8);
3rd gill openings, 1020(22.0, 3 3.0); 4th gill open-
ings, 1150 (25.8, 37.2); 5th gill openings (head
length), 1 180 (26.5, 38.2); pectoral origins, 1 1 10
(24.9, 35.9); pelvic origins, 2270 (50.9, 73.5); 1st
dorsal origin, 1540 (34.5, 49.8); 2nd dorsal ori-
gin, 2530 (56.7; 81.9); anal origin, 2830 (63.5,
91.6); vent, 2295(51.5, 74.3).

Distance between: vent and caudal tip, 2165
(48.5, 70. 1); 1st and 2nd dorsal origins, 625 (14.0,
20.3); 1st and 2nd dorsal bases, 590(13.2, 19.1);
2nd dorsal and upper caudal origins, 428 (9.6,
13.9); 2nd dorsal base and upper caudal origin,
395 (8.9, 12.8); pectoral and pelvic origins, 510
(11.4, 16.5); pelvic and anal bases, 330(7.4, 10.7);
anal and lower caudal origins, 315 (7.1, 10.2);
anal base and lower caudal origin, 230 (5.2, 7.4).

Eyes (palpebral apertures or fleshy orbits):
length, 56 (1.3, 108); height, 54 (1.2, 1.7); width
across anterior corners (interorbital), 370 (8.3,
12.0); eyeball diameter, 84 (1.9, 2.7).

Nostrils: width, 30 (.07, 1 .0); internarial space,
340 (7.6, 8.8).

Spiracles: diameter, 6 (0. 1, 0.2); space between
spiracles and eyes, 176 (3.9, 5.7).

Mouth (jaws in retracted position): length, 273
(6.1, 8.8); width, 827 (18.5, 26.8); width across
outer edges of jaws, 1025 (23.0, 33.2); length of
lower jaw, 820(18.4,26.5).

Gill opening widths (heights): 1st, 265 (5.9,
8.6); 2nd, 258 (5.8, 8.4); 3rd, 264 (5.9, 8.5); 4th,
256 (5.7, 8.3); 5th 234 (5.2, 7.6).

Head height: at spiracles, 500 (1 1.2, 16.2); at
1st gill openings, 625 (14.0, 20.2); at 5th gill
openings, 630 (14.1, 20.4).

Trunk height: at 1st dorsal origin, 640 (14.3,
20. 7); at pelvic origins, 515(11.5, 16. 7); at pelvic
insertions, 440 (9.9, 14.2).

Girth: at 1st dorsal origin, 1800 (40.4, 58.2);
at 2nd dorsal origin, 1 140 (25.6, 36.9).

Caudal peduncle height: at 2nd dorsal inser-
tion, 341 (7.6, 1 1.0); at upper caudal origin, 237
(5.3, 7.7).

Caudal peduncle width: at 2nd insertion, 146
(3.3, 4.7); at upper caudal origin, 109 (2.4, 3.5).

Pectoral fins, length of: anterior margin, 837
(18.8, 27.1); posterior margin, 615 (13.8, 19.9);
base, 262 (5.9, 8.5); origin to free rear tip, 453
(10.1, 14.7); inner margin, 190 (4.3, 6.1).

Pelvic fins, length of: anterior margin, 264 (5.9,
8.5); posterior margin, 181 (4.1, 5.9); base, 207
(4.6, 6.7); origin to free rear tip, 245 (5.5, 7.9);
inner margin, 38 (0.8, 1.2); height, 255 (5.7, 8.3);
origin to rear tip of clasper, 575 (12.9, 18.6).

Claspers: inner length from vent to tip, 550
(12.3, 17.8); outer length from clasper base to
tip, 355 (8.0, 11.5); width at outer pelvic base,
47(1.1, 1.5).

1st dorsal fin, length of: anterior margin, 415
(9.3, 13.4); posterior margin, 265 (5.9, 8.6); base,
404(9.1, 13.1); inner margin, 82 (1.8, 2.7); height,
226(5.1,7.3).

2nd dorsal fin, length of: anterior margin, 198
(4.4, 6.4); posterior margin, 158 (3.5, 5.1); base,
191 (4.3, 6.2); inner margin, 80 (1.8, 2.6); height,
104 (2.3, 3.4).

Anal fin, length of: anterior margin, 196 (4.4,
6.3); posterior margin, 80 (1.8, 2.6); base, 159

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. Vol. 43, No. 8

FIGURE 5. (a) Schematic head drawing ofMegachasma pe-
lagios with jaws protruded, (b) Photograph of fresh shark with
jaws protruded. (Upper arrow points to anteriormost edge of
neurocranium; lower arrow to upper jaw and teeth.)

(3.6, 5.1); inner margin, 67 (1.5, 2.2); height, 78
(1.7,2.5).

Caudal fin, length of: dorsal margin, 1443 (32.3,
46.7); preventral margin, 625 (14.0, 20.2); lower
postventral margin, 377 (8.5, 12.2); upper post-
ventral margin, 1220 (27.4, 39.5); subterminal
margin, 57 (1.3, 1.8); terminal margin, 96 (2.2,
3.1); terminal lobe or sector, 1 39 (3. 1 , 4.5); width
of dorsal lobe at postventral notch, 471 (10.6,
1 5.2); width of ventral lobe at postventral notch,
273(6.1,8.8).

Intestinal valve: length, 690 (15.5, 22.3); di-
ameter, 145 (3.3, 4.7); thickness of broadest an-
terior ring, 25 (0.6, 0.8).

DESCRIPTION (based on the holotype and only
known specimen). — Head length from snout tip
to 5th gill openings, 26% of total length and 1.6
times distance between pectoral and pelvic fin
bases. Head broad, cylindrical, and approxi-
mately circular in transverse section at eyes, but
somewhat laterally expanded and oval in section

over jaws when jaws are retracted; not depressed.
Outline of head in lateral view nearly straight
dorsally, except for bluntly convex snout, strong-
ly convex ventrally along edges of lower jaws and
nearly straight beneath gills; in dorsoventral view,
anteriorly rounded and convex and tapering pos-
teriorly to gills. Snout length from tip to edge of
mouth about 12.5 times in mouth width. Snout
broadly rounded in dorsal view, with lateral mar-
gin slightly indented anterior to nostrils; in lateral
view, convex dorsally and concave ventrally to
fit the front of the retracted upper jaw (Fig. 5).
External eye opening (palpebral aperture) or fleshy
orbit without anterior or posterior notches, length
about 2 1 times in head length. Irises of eyes black,
nearly filling orbits. Eyeballs large, diameter 14
times in head length. Spiracles small, their lengths
about '/io orbit length, located about 3 orbit
lengths behind eyes and about opposite ventral
margins of eyes. Gill openings of nearly equal
length, the longest ( 1 st and 3rd) about 4.5 in head
length and 4.7 times eye length, the smallest (5th)
about Vio length of longest. Edges of gill openings
nearly straight, not incised, and with filaments
not exposed when jaws are retracted. Gill open-
ings with upper ends falling below level of eyes,
and midheight of head at gill openings. Internal
gill openings with numerous gill-raker papillae
arranged in about 4 rows on their anterior and
posterior edges, including both anterior and pos-
terior edges of 1 st gill cavity between hyoid and
1 st branchial arches and posterior edge of 5th
gill cavity on anterior edge of 5th gill arch. Gill-
raker papillae small, about 10-15 mm long,
densely packed, slender, tapering to blunt point,
arranged with tips pointing anteromedially into
pharynx, with thick epidermis and dermis cov-
ering hyaline cartilage core layered with flat-
tened, imbricated denticles (Fig. 6). Nostrils with
large lateral incurrent aperture, anterior nasal flap
with an undulated, truncated posterior edge, and
low keel on dorsal surface, but no distinct meso-
narial flap, small medial excurrent aperture with
low posterior nasal flap on its rim. Nostrils lateral
to mouth edge and 2.4 times closer to snout tip
than to eyes. Nostril width 1.8 in orbit length,
8.8 times in longest gill opening. Inner labial
grooves at mouth corners on both upper and
lower jaws just lateral to dental bands and medial
to vertical fold of skin sheathing adductor man-
dibulae muscles. Mouth width when jaws are
retracted about 1 .4 in head length; mouth length

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

FIGURE 6. Gill-raker papillae of Megachasma pelagios. (a) Drawing of a group of gill-raker papillae, (b) Scanning electron
micrograph of the tip of a single papilla, showing the closely imbricated denticles (20X magnification), (c) Scanning electron
micrograph of denticles from b, at higher magnification (5 1 X).

about 3 times in width. A broad fold of skin
forming a deep pocket on dorsal surface of upper
jaws below snout, and a vertical fold of skin en-
closing anterior edges of adductor mandibulae
muscles at each mouth corner. Tips of upper jaws
can extend at least 6 orbit lengths in front of
snout tip, with mouth corners passing anterior

to eyes. Tongue extremely large, broadly round-
ed and thick, enclosing greatly enlarged basihy-
oid cartilage; tongue almost entirely filling mouth
cavity when jaws are closed. Deep pocket under
front of tongue, freeing it anteroventrally; pocket
about 4 orbit lengths deep from anterior tongue
edge to its basal attachment to mouth. Maxillary

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8

FIGURE 7. Illustrations of (a) upper symphysis and (c) lower symphysis of Megachasma pelagios, showing bare, toothless
patches and mesial ends of dental bands, (b) Close-up of rows of teeth from upper jaw.

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

101

DRL

FIGURE 8. Illustration of a lower tooth of Megachasma
pelagios from about midlength on the lower left dental band
in (A) labial; (B) lingual; (Q distal; and (D) basal views. Ab-
breviations: AS, attachment surface of root; BG, basal groove;
BL, basal ledge; CLGF, centrolingual foramen; CR, crown; CU,
cusp; DCE, distal cutting edge; DRL, distal root lobe; DSH,
distal shoulder; LAP, labial foramina; LGP, lingual protuber-
ance; MRL, mesial root lobe; MSH, mesial shoulder; NK, neck;
RT, root; TG, transverse groove. Drawn by L. J. V. Compagno.

valve of upper jaw arcuate and relatively narrow,
width less than an eye diameter. No enlarged
buccal papillae in mouth cavity, but with scat-
tered circular organs of undetermined function
on tongue and mouth.

Teeth very small and relatively numerous, in
56 rows in left upper, 59 rows in left lower, 52
rows in right upper, and 69 rows in right lower
jaw halves, or 56-52/59-69; total tooth-row
counts 108/128. Teeth not arranged in diagonal
files. Symphyseal toothless space about 4 orbit
diameters wide in upper jaw and less than one
eye-length wide in lower jaw (Fig. 7). Dental bands
of upper and lower jaws show strong gradient
monognathic heterodonty; starting from small
teeth at symphysis, teeth increase in size to about
10 mm high in about 10 tooth rows distal to
symphysis, then begin to gradually decrease in
size and increase in width relative to height to
distal ends of dental bands. Teeth (Fig. 8) have
no cusplets, narrow crown shoulders, partial
transverse groove on linguobasal attachment
surface of root, large centrolingual foramen, and
scattered labial foramina below basal ledge. A
sectioned tooth (Fig. 9) shows thick osteodentine

FIGURE 9. Diagrammatic sagittal section of a lower tooth
of Megachasma pelagios from about midlength on the lower
left dental band, lingual protuberance broken off. Abbrevia-
tions: CR, crown; CU, cusp; END + ORD, enameloid + or-
thodentine; OSD, osteodentine; RT, root. The teeth of M.
pelagios are of the "osteodont" type, with a crown having a
core of osteodentine and no pulp cavity or canal. Drawn by
L. J. V. Compagno.

core in crown, surrounded by relatively thin lay-
ers of pallial orthodentine and enameloid, and
no pulp canal or cavity; crown osteodentine con-
tinuous with that of root, which forms its sole
component.

Body stout, trunk circular or vertically oval in
section at first dorsal base. Length of head and
trunk from snout tip to vent 50% of total length.
Trunk relatively short, length from 5th gill open-
ing to vent 1 . 1 times head length. No predorsal,
interdorsal, or postdorsal ridges on midline of
back and precaudal lobe; no lateral ridges on
body. Precaudal lobe from vent to upper caudal
origin short, 19.2% of total length. Height of cau-
dal peduncle at insertion of second dorsal 2.3
times its width there and 1.2 times in distance
from insertion of second dorsal to upper caudal
origin; height of caudal peduncle at upper caudal
origin 2.2 times its width there and 1.7 times in
distance from insertion of second dorsal to upper
caudal origin. Upper precaudal pit not transverse
and crescentic.

Dermal denticles on body very small and flat-
tened, giving skin a smooth texture. Denticles on
sides of trunk below first dorsal fin (lateral trunk
denticles) loosely spaced, not closely imbricated
(Fig. 1 0), with a strong medial ridge and a pair
of strong lateral ridges running entire length of
crown, strong medial cusp, but with lateral cusps
absent or hardly developed. Denticles on dorsal
surfaces of pelvic fins (Fig. 1 1) similar to lateral

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8

FIGURE 10. Scanning electron micrographs of dermal den-
ticles of Megachasma pelagios. (top) Lateral trunk denticles
from just below the base of the first dorsal fin (60X magnifi-
cation), (bottom) Denticles from the surface of the tongue (67x).

trunk denticles, except for having lower ridges
and being closely imbricated. Denticles from
tongue are wide-spaced but broader and more
transversely oval than lateral trunk denticles.
Distal webs of upper surfaces of pectoral and
pelvic fins, and dorsal caudal lobe, have con-
spicuous dark wavy lines, often parallel, which
are channels of bare skin between areas of den-
ticulate skin (Fig. 11).

Pectoral fins moderately broad basally but dis-
tally elongated, tapering, falcate, and broad-
tipped. Anterior margins of pectoral fins mod-
erately convex, apices broadly angular, posterior
margins slightly convex, and free rear tips and
inner margins smoothly rounded and broadly
convex. Length of pectoral fin from origin to free
rear tip 1.9 times in its anterior margin length.

Apex of pectoral posterior to its free rear tip when
fin is appressed to body.

Pectoral fin skeleton with all radials except last
5 on metapterygium greatly elongated, with
broad, flattened tips. Radials with numerous seg-
ments, the longest with 10; distalmost segments
elongated but only about '/3 length of each radial.
Pectoral fin propterygium supporting one radial,
mesopterygium with 5 radials, metapterygium
with 8 radials on basal segment and 8 on axis.
Propterygium small and slightly elongated dis-
tally. Mesopterygium moderately elongated dis-
tally, fairly broad and wedge-shaped with radials
inserted on distal end at an angle to axis of elon-
gation. Metapterygium diagonally elongated
across fin base with radials inserted at an angle
to long axis. Metapterygial axis of 5 segments,
about % as long as basal metapterygium. Basal
and radial cartilages of pectoral fins not highly
calcified; fins rather flexible, despite having ple-
sodic skeletons.

Pelvic fins with anterior margin slightly con-
cave anteriorly but convex posteriorly, apex very
narrowly rounded, and inner margins slightly
concave. Inner margins, posterior margins, and
free rear tips of pelvics forming broad triangle.

Claspers relatively slender, width at base 7.6
times outer length from pelvic bases to tips, inner
length from vent to tip 1 2.3 percent of total length.
Rear tips of claspers reaching almost to midbase
of anal fin when claspers are horizontal. Clasper
tip elongated, forming a very narrow, slender
process (Fig. 1 1), glans anterior to elongated tip
slightly spatulate and flattened, shaft cylindrical.
Clasper groove open, with edges not fused dor-
sally; no pseudopera or lateral clasper groove and
fold. Small, sharp-tipped, hardened clasper spur
on ventral lobe, lateral to groove. Large, large-
mouthed, prominent pseudosiphon on the dorsal
clasper lobe.

First dorsal fin with anterior margin slightly
concave anteriorly and convex posteriorly, pos-
terior margin nearly straight, free rear tip acute
and slightly attenuated, and inner margin slightly
concave. Origin about opposite or slightly pos-
terior to pectoral fin insertions, midpoint of dor-
sal base about 2.6 times closer to pectoral inser-
tions than pelvic origins, dorsal fin insertion
anterior to pelvic origins by about 0.8 times first
dorsal base, and free rear tip about 2.9 times
dorsal inner margin anterior to pelvic origins.
Posterior margin slanting posteroventrally from
dorsal apex, insertion well posterior to level of

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

103

FIGURE 11. Right pelvic fin and clasper of Megachasma pelagios, showing channels of naked skin on dorsal surface of pelvic
fin. (a) Close-up photograph of the channels, (b) Scanning electron micrograph of denticles from the anterior edge of the pelvic
fin (54X magnification), (c) Drawing of the pelvic fin and clasper.

dorsal apex. Base 1.5 times in interdorsal space
and 3.6 times in dorsal caudal margin, height 1 .8
times in base, and inner margin 2.8 times in
height.

Second dorsal fin low, height 0.46 times first
dorsal height, base 0.47 times first dorsal base.

Anterior margin nearly straight, apex narrowly
rounded, posterior margin slightly concave, free
rear tip angular and attenuated, and inner margin
slightly concave. Free rear tip about over anal
fin origin. Posterior margin of second dorsal
slanted posteroventrally from apex, insertion

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8

Sagittal Section

CCA

VRA

NOS
Transverse Section

NAR

FIGURE 12. Monospondylous precaudal vertebrae of Me-
gachasma pelagios, in (top) sagittal section, and (bottom) trans-
verse section. Abbreviations: Sagittal section: CCA, central
canal; CCE, cartilaginous centrum; NO, notochord; NOS, no-
tochordal sheaths; VRA, vestigial radii. Transverse section:
CCA and VRA, as above; BD, basidorsal; BV, basiventral; DI,
dorsal intermedial; LIN, lateral intermedial; NAR, neural arch;
VI, ventral intermedial. (From drawings by L. J. V. Com-
pagno.)

posterior to apex. Base 2. 1 times in distance from
insertion to upper caudal origin, height 1 .8 times
base, and inner margin 1 .2 times height.

Anal fin low, height 0.8 times second dorsal
height, base length 0.8 second dorsal base. An-

terior margin concave anteriorly but convex pos-
teriorly, apex broadly rounded, posterior margin
moderately concave or notched, free rear to acute
and attentuated, and inner margin slightly con-
cave. Posterior margin of anal fin slanted pos-
terodorsally from apex, with anal apex just below
insertion. Base 1 .4 times in distance from inser-
tion to lower caudal origin, height 2.0 times in
base, and inner margin 1.2 times in fin height.

Caudal fin relatively asymmetrical, with ba-
sally broad dorsal lobe, and short terminal lobe.
Length of dorsal margin 2. 1 times in precaudal
length, of preventral caudal margin 2.3 times in
dorsal caudal margin, and of terminal lobe from
caudal tip to subterminal notch about 10.3 times
in dorsal caudal margin. Dorsal caudal margin
slightly but continuously convex in lateral view,
preventral margin almost straight dorsally but
becoming more convex ventrally. Tip of ventral
caudal lobe broadly angular, lower and upper
postventral margins slightly convex, notch be-
tween postventral margins broadly angular, sub-
terminal notch shallowly concave, subterminal
margin slightly concave, and terminal margin
slightly convex. Subterminal margin length 0.6
times terminal margin length. Ventral lobe of
caudal fin aplesodic, not supported by hypural
radials but by ceratotrichia and connective tissue
only.

Vertebrae (Fig. 12) examined from beneath
first dorsal fin (monospondylous precaudal ver-
tebrae) and at base of caudal fin (diplospondylous
caudal vertebrae). These found to have ex-
tremely reduced calcification, both of the pri-
mary double cone of vertebral centra (which is
almost entirely formed of uncalcified cartilage
and connective tissue in M. pelagios), and of
intermedial areas between basidorsals and bas-
iventrals. Vertebral centra consist of biconic or
bioconcave discs of cartilage, separated by broad
bands of unchondrified notochordal sheath and
spherical cavities containing notochordal tissue.
Calcification in monospondylous precaudal cen-
tra restricted to some irregular calcification on
lateral centrum body, a layer on ventral part of
neural canal, a layer on midventral groove on
underside of centrum, and paired thin zones partly
bounding intermedial areas between basals, in-
cluding 2 dorsals, 2 ventrals, and 2 pairs of lat-
erals. These intermedial calcifications resemble
radii of other lamnoids, but differ in being only
partially developed across intermedial areas and
in not forming discrete longitudinal plates. These

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

105

Dorsal View

Ventral View

Lateral View

FIGURE 13. Neurocranium of Megachasma pelagios, in dorsal, ventral, and lateral views. Abbreviations: Dorsal view: AF,
anterior fontanelle; CR, cranial roof; LR, lateral rostral cartilage; MR, medial rostral cartilage; NA, nasal aperture; OR, opisthotic
ridge; OT, otic capsule; PF, parietal fossa; PR, preorbital process; PRF, profundus foramen; PT, postorbital process; RN, rostral
node; SC, supraorbital crest; SF, supraorbital fenestra; SRI, sphenopterotic ridge. Ventral view: LR, MR, and OT as above; AP,
articular pit; BP, basal plate; ECF, ectethmoid foramen; HF, hyomandibular facet; ICF, internal carotid foramen; INS, internasal
septum; NC, nasal capsule; SC, suborbital crest; SS, suborbital shelf; STF, stapedial fenestra. Lateral view: O, orbit; ORF, orbital
fissure; SC, suborbital crest; SCA, sphenopterotic capsule; all others as above. (From drawings by L. J. V. Compagno.)

intermedial calcified zones interpreted as rep-
resenting vestigial radii, greatly reduced in Me-
gachasma but probably well developed in its pre-
cursors. Basal caudal centra similar to
monospondylous precaudal centra, except for
having intermedial calcifications even more re-
duced to a set of dorsal and ventral pairs only.
The poorly calcified vertebral centra of Mega-
chasma recall the septate vertebral columns of
large species ofSomniosus (subgenus Somniosus,
for S. pacificus and S. microcephalus) and some
other squaloids (see Compagno 1977), with re-
duction of form and calcification of centra and
hypertrophy of notochordal tissue in between
centra. The lamnoids Mitsukurina and Pseudo-
carcharias have extremely simple centra with
double cones and radii reduced to 8 slightly
branched plates (2 bounding each intermedial
area), but Megachasma goes far beyond these

genera in reduction of its centra, in calcification,
and in intrusion of notochordal tissue. Mitsu-
kurina and Pseudocarcharias retain normal, close-
set double cones, despite their simple radii.

Neurocranium (Fig. 1 3) dissected on one side
only, and reconstructed bilaterally. Cranium rel-
atively large, extremely broad and moderately
flat; nasobasal length (from base of medial rostral
cartilage to occipital condyles) about 8.9 percent
total length and 12.8 percent precaudal length;
greatest width of cranium across preorbital pro-
cesses about equal to nasobasal length, and great-
est height from cranial roof to ventral edges of
suborbital shelves 0.4 times in nasobasal length
and greatest cranial width. Rostrum relatively
short but very broad, length of medial rostral
cartilage from its base to anterior edge of rostral
node about 26 percent nasobasal length; width
across outer bases of lateral rostral cartilages 2.2

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8

Neurocranium
SC 0 OP

FIGURE 14. Jaw suspension of Megachasma pelagios, in lateral view, jaws retracted. Abbreviations: H, hyomandibular; MC,
Meckel's cartilage; OP, orbital process; PP, palatine process; PQ, palatoquadrate; QP, quadrate process; VC, vertebral column;
all others as in Figure 12. (From drawings by L. J. V. Compagno.)

times length of medial rostral cartilage. Entire
ethmoid region of cranium, including rostrum,
nasal capsules, and internasal septum, and the
anterior basal plate molded dorsally around en-
larged palatine processes of palatoquadrates (Fig.
14). Lateral rostral cartilages with broad bases
that cover entire anterior surfaces of nasal cap-
sules; diagonally compressed from dorsomedial
to ventrolateral, and extending anteromedially
as tapering triangular bars to meet rostral node
separately on either side. Medial rostral cartilage
and its base on internasal septum deflected up-
ward over symphysis of palatoquadrates (jaws in
retracted position), so that base originates at a
level slightly above lateral rostral cartilages and
the moderately depressed, narrow, barlike shaft
arches anterodorsally and anteroventrally to ros-
tral node. Rostral node a simple, depressed,
narrow plate, not vertically or horizontally fe-
nestrated, anteriorly expanded, vertically com-
pressed, nor with rostral appendices.

Nasal capsules extraordinarily modified, high-
ly compressed, platelike, wedge-shaped struc-
tures with nasal fenestra mainly on their lateral
faces. Plane of compression of nasal capsules
congruent with large-based lateral rostral carti-

lages, together forming a lateral wall to expansion
cavity enclosing palatine processes. Ectethmoid
foramen present on dorsomedial surface of each
nasal capsule. Large subethmoid fossa on ventral
surface of depressed, laterally expanded inter-
nasal septum, extending anteriorly beneath ros-
trum and medially to nasal capsules, and pos-
terolaterally to merge on either side with large
orbital process cavity in basal plate. Foramina
for nasal canals laterally situated in cranial cavity
(not anterolateral), with canals running antero-
laterally to nasal capsules.

Basal plate very broad, width across orbital
notches about 68 percent nasobasal length,
broadly arched over rear ends of palatoquadrate
palatine processes (when retracted) but relatively
flat posterior to internal carotid foramina. Entire
ventral surface of suborbital shelves, basal plate,
and internasal septum padded with thick, soft,
spongy connective tissue, probably to cushion it
from palatoquadrates. Basal plate with pair of
internal carotid foramina located about 59 per-
cent nasobasal length behind medial rostral car-
tilage, separated by a convex space with width
80 percent nasobasal length and 1 . 1 times in dis-
tance between internal carotid foramina and sta-

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

107

pedial fenestrae. Stapedial fenestrae small, width
about 3 percent nasobasal length, apertures about
1.6 times closer to internal carotid formina than
to lateral edges of suborbital shelves. Stapedial
fenestrae apparently without greatly convoluted
arteries or a rete mirabile elaborated from effer-
ent spiracular arteries. Basal plate nearly hori-
zontal posteriorly, without medial keels.

Orbits nearly circular in lateral view, with large
optic nerve foramen slightly dorsal to its center.
Orbits moderately large, with horizontal diam-
eters about 43 percent of nasobasal length. Su-
praorbital crests broad, not fenestrate basally,
only moderately concave in dorsal view. Preor-
bital processes not strongly exserted from su-
praorbital crests, and extending ventrally to pos-
terior edges of nasal capsules. Small preorbital
canal fenestra for superficial ophthalmic nerves
present between broad preorbital process and
cranial roof on each side; profundus nerve fo-
ramen just mesial to fenestra. Postorbital pro-
cesses ventrally produced almost to level of optic
nerve foramen, bifurcate distally. Foramina of
orbital wall not examined in detail but including
foramina for superficial ophthalmic nerve an-
terior cerebral veins, optic nerve, and large, deep,
trigeminofacialis chamber or orbital fissure. Sub-
orbital shelves nearly vertical, large, thick basally
but distally thin, arcuate, and with sides nearly
parallel in ventral view.

Otic capsules large and subquadrate, with
lengths about 36 percent nasobasal length and
width about 82 percent nasobasal length. Hyo-
mandibular facets huge, ventromedially incised,
and broadly arcuate, extending along entire length
of otic capsules from otic processes anteriorly to
partway onto bases of suborbital shelves, but not
exserted posteriorly from occiput. Hyomandib-
ular nerve foramina just below opisthotic ridges
and about midway along their lengths on otic
capsules. Sphenopterotic ridges arching postero-
medially in dorsal view, ending posteriorly in a
bluntly rounded corner. Opisthotic ridges on
dorsal surface of hyomandibular facets low and
curved posteroventrally. Occiput flat and not
exserted rearwards, with glossopharyngeal and
vagus nerve foramina.

Jaws (Figs. 5, 14) poorly calcified; length of
palatoquadrates about 16 percent total length,
Meckel's cartilages 1 8.4 percent total length. Pal-
atine processes of palatoquadrates articulating at
symphysis and extending for about % of pala-

toquadrate length to orbital processes. Meckel's
cartilages huge, ventrally arcuate, dorsally nearly
straight, thick, and compressed, with long pos-
terior extensions from their mandibular articu-
lations with palatoquadrates. Meckel's cartilages
articulating closely at mandibular symphysis.

Manipulation of the jaws of the fresh-caught
Megachasma pelagios suggested that the jaws are
highly protrusible, but not necessarily as a mech-
anism to quickly eject them outward to capture
prey, as in some other lamnoids (most notably
Mitsukurind), nor to bring the upper teeth to bear
on prey items, as in Carcharodon carcharias. The
jaws may be protruded forward and outward to
expand the mouth aperture and form a hoop-net
for capturing plankters, though we do not know
the exact shape of the jaws deployed in this con-
figuration without photographic documentation
of a live M. pelagios feeding. The basking shark
is able to deploy its much slimmer jaws almost
in a circle while feeding and has been photo-
graphed many times with jaws expanded (but not
protruded); however, the exact shape of the mouth
opening in a living, feeding basking shark would
be somewhat difficult to work out from a dead,
preserved specimen. The jaw structure of M. pe-
lagios suggests that the jaws move downward,
anteriorly, and outward at the mouth corners,
and the distal ends of the hyomandibulae swing
anterolateroventrally as protrusion occurs. The
mechanism of jaw protrusion is poorly under-
stood with the limited dissection possible during
preparation of this description (the desire to limit
damage to the specimen prior to making a cast
of it prohibited a thorough investigation of the
jaw mechanism and the hyobranchial skeleton
and musculature), but the large, straplike, diag-
onal preorbitalis muscles may help to pull the
jaws forward.

The jaw musculature was not investigated in
detail, but sufficient information was collected
to determine that the jaw muscles are similar to
those in other lamnoids. Levator palatoquadrati
muscle simple, originating on Sphenopterotic
ridges of otic capsules and running posteroven-
trally to insert on quadrate processes of palato-
quadrates. Adductor mandibulae muscles mod-
erately large but small and weak compared to the
huge jaws, and limited anteriorly by mouth cor-
ners. Levator hyomandibuli muscles broad and
relatively large.

The viscera were not examined in detail, ex-

108

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8

FIGURE 15. Drawing of the valvular intestine of Megachasma pelagios with the dorsolateral quadrant removed to show the
ring valves with highly fimbricated edges (anterior to the right).

cept to note that the liver is relatively small
(though very oily) and that the valvular intestine
is an elongated ring- valve type rather similar to
that of other lamnoids (Fig. 15). Each ring has a
densely fimbriated surface, apparently to in-
crease the absorptive surface, with a maximum
thickness of 23-25 mm at the pyloric end of the
intestine but becoming increasingly thin towards
the rectum.

Color. When preserved, dorsal surface of head,
trunk and tail, dorsal surfaces of pectoral and
pelvic fins, dorsal fins, center of anal fin, and
caudal fin dark gray to blue-black; sides of body
lighter, underside of body and fins light gray,
except for underside of head and lower jaw which
are dark gray and mottled with black, undersur-
face of pectoral fins which have an abruptly black
anterior margin about an eye diameter wide, and
underside of pelvic fins with a dusky anterior
margin. Tips and posterior margins of dorsal sur-
faces of pectoral and pelvic fins abruptly white,
as are posterior margins of dorsal and anal fins
and postventral caudal margins. Tissue of oral
cavity and tongue blackish in preservative, but
with oral lining silvery when fresh.

FEEDING HABITS AND BIOLOGY

The stomach contents were a thick reddish
soup abundantly stocked with the euphausiid
shrimp Thysanopoda pectinata, a species attain-
ing a median length of 31 mm (Hu 1978). Ac-
cording to Hu (1978), T. pectinata off the west
coast of Oahu (21°15'-20'N, 158°15'-30'W)
shows a moderate day and night migration pat-
tern. During the day most are caught between
350 and 750 m depth, with some ranging up to
300 m and down to 1100 m, but at night the

bulk are between 150 and 500 m depth, with
some up to 75 m and down to 525 m. Apparently,
when captured, Megachasma pelagios would
have been in the upper depths (165 m) where
these euphausiids are commonest at night, and
quite possibly might have been feeding on them
when it became entangled in the parachute.

The megamouth shark unites an eclectic com-
bination of habitus characters that (along with
its apparent epipelagic habitat and filter-feeding
habits) suggests an unusual mode of life. Deep-
water epibenthic and epipelagic sharks often show
a decrease in specific gravity and increase in hy-
drostatic support by the enlargement of their ab-
dominal cavity and liver volume to produce a
large, oily, hepatic "float." M. pelagios, in con-
trast, has reduced specific gravity in the form of
extremely poor calcification; a soft, almost en-
tirely hyaline cartilage skeleton; very soft, loose
skin; and flabby, loose connective tissue and
muscles. These features, and its soft, rubbery pre-
caudal fins; lack of a keel on the caudal peduncle,
weak precaudal pit; lack of dorsal caudal ripples;
and highly flexible, asymmetric caudal fin suggest
that M. pelagios is a slow, weak swimmer.

It is interesting to compare M. pelagios with
the other two species of large, filter-feeding sharks:
the basking shark and the whale shark. The bask-
ing shark is the only lamnoid filter-feeder besides
megamouth, but in contrast has many adapta-
tions for a higher activity level and sustained
powerful swimming, including a strongly calci-
fied skeleton, firm muscles, stiff fins, dense skin,
and tough connective tissue; a huge, oily liver
and elongated body cavity; a more fusiform body,
lunate caudal fin, strong caudal keels and pre-
caudal pits, and huge gill openings. The filter

TAYLOR, CAMPAGNO & STRUHSAKER: MEGAMOUTH-A NEW SHARK

109

apparatus of Cetorhinus, with its vast gill cavities
and slender, smooth, streamlined gill-raker den-
ticles, is clearly adapted for a higher rate of water
flow than is possible with the smaller gill cavities,
more restricted internal gill apertures, and less
streamlined gill-raker papillae of Megachasma.
The basking shark is a slow but strong swimmer,
which has often been observed and photo-
graphed while feeding at or near the surface with
its mouth distended to form a circular scoop.
Although its mouth is relatively smaller than that
of megamouth, the basking shark is probably a
much more efficient dynamic filterer because of
its stronger swimming abilities and high-flow ni-
ter apparatus. The prey of the basking shark is
far smaller than what is known for megamouth,
consisting entirely of microscopic crustaceans
(especially copepods). The basking shark prefers
cool to cold coastal waters rich in nutrients and
plankton.

The whale shark resembles the basking shark
in its strong swimming adaptations, except that
it has a less fusiform body, flattened anteriorly;
a shorter body cavity and much smaller liver;
and much smaller external gill openings (but larg-
er than those of megamouth). The filter appa-
ratus of Rhiniodon differs from that of Mega-
chasma and Cetorhinus in not being confined to
the margins of the internal gill openings; instead,
the gill filter elements of Rhiniodon cross and bar
these openings. They are compressed, triangular,
cartilage-cored, connective-tissue-covered, par-
allel plates that transversely bridge the internal
gill openings and connect adjacent holobranchs.
The plates have highly lobulated pharyngeal
margins that form an interconnected network, or
dense filter grid, and are divided into paired dor-
sal and ventral groups of plates or screens over
each internal gill opening. The dense screens of
Rhiniodon are obviously efficient filters, but are
incapable of sustaining a high flow of water
through them. However, this filter apparatus,
combined with a broad but very short, transverse
mouth; very long, broad, low pharynx, and rel-
atively small gill openings apparently adapts the
whale shark to a combination of suction feeding
(as in Ginglymostoma and other orectoloboids)
and filter-feeding not found in Megachasma and
Cetorhinus. The bellowslike pharynx and filter
screens of the whale shark may provide it with
a more versatile feeding apparatus than in Ceto-
rhinus (and presumably Megachasma) by allow-
ing it to suck in and filter out a wide variety of

prey animals, independent of the shark's forward
movement. Although the whale shark can ingest
small crustaceans, it also eats squid and com-
monly takes small schooling fishes such as an-
chovies and sardines, and even small albacore
and tuna (Bigelow and Schroeder 1 948). It is not
known whether Rhiniodon can filter out crus-
tacean prey as small as the copepods favored by
Cetorhinus, but almost certainly the euphausiids
eaten by Megachasma are in the prey-size range
of the whale shark, which is a warm-temperature
to tropical, coastal to oceanic, slow but strong-
swimming shark, often seen basking or cruising
at the surface and feeding on schools of fishes. It
often positions itself vertically beneath a school
of prey, unlike the horizontal attitude Cetorhinus
maintains while feeding at the surface.

The soft, flabby body and fins, low-flow bran-
chial filter apparatus, and small gill openings sug-
gest that Megachasma is less active and possibly
a less efficient filter-feeder than Cetorhinus or
Rhiniodon. Nevertheless, this species has a spe-
cialized, presumably efficient mechanism for
capturing small oceanic animals in its oversized
jaws which are enlarged to increase the diameter
of its "net" and thickened to provide adequate
support from its rubbery hyaline cartilage. The
greatly distensible mouth and pharynx, closely
packed gill-raker papillae, and large tongue prob-
ably help to expel water from the pharynx when
it closes its mouth. Megachasma can be imag-
ined as slowly swimming through schools of eu-
phausiid shrimp and possibly other prey with
jaws widely opened, occasionally closing its
mouth and contracting its pharynx to expel water
and concentrate its prey before swallowing it.

Inspection of the mouth of megamouth 24 hr
after capture revealed a bright silvery lining
punctuated by small circular porelike structures.
At the time it was speculated that these might
be bioluminescent organs, but we have no evi-
dence of this. Histological sections of mouthlin-
ing were made but were problematical because
of the deteriorated state of the tissue.

That Megachasma may not be a more active
filter-feeder such as Cetorhinus or Rhiniodon may
be related to its tropical deepwater oceanic hab-
itat, which has a relative paucity of nutrients and
prey in comparison to the cool coastal surface
waters favored by Cetorhinus and the tropical
coastal waters preferred by Rhiniodon. Various
mesopelagic teleosts have reduced skeletal and
other tissues as adaptations to a nutrient-poor

110

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 8

environment, and Megachasma may be similar-
ly limited to a reduced level of tissue develop-
ment and hence a low activity level for a filter-
feeding shark, far less than is possible in the
habitats frequented by Cetorhinus and Rhinio-
don.

Two distinctive scars, one on the throat,
another behind the right pectoral fin, suggest that
megamouth may be the only known selachian
victim of Isistius brasiliensis, the "cookie-cut-
ter" shark, that is believed responsible for similar
marks found on tuna, porpoise, and billfish caught
in Hawaiian waters (Jones 1971). The soft skin
and midwater habitat of megamouth may make
it vulnerable to Isistius attacks.

Megachasma pelagios, itself the representative
of a new family of sharks, is the host of a new
family of tapeworms Mixodigmatidae (order
Trypanorhynchida), described by Dailey and
Vogelbein (1982) for the new genus and species
Mixodigma leptaleum. These parasitic tape-
worms from the valvular intestine presented
taxonomic problems over placement in existing
trypanorhynch families comparable to the diffi-
culties encountered in attempting to place Me-
gachasma pelagios in an existing lamnoid shark
family.

ACKNOWLEDGMENTS

Many people contributed to the study of this
interesting animal. We acknowledge Lt. Linda
Hubble, John Hobbs, and Rick Kahakini for their
recovery of the animal and their recognition of
the find; Hawaiian Tuna Packers, National Ma-
rine Fisheries Services Honolulu dockside staff,
Phillip Motta, Bruce Carlson, Captain Gary Naf-
tel and the crew of the R.V. Easy Rider, Mathew
James, Les Matsuura, Marge Awai, and Arnold
Suzumoto for curatorial assistance; Dr. Arthur
Popper, Dr. James Margolis, and Karen Margolis
for preparation of scanning electron micro-
graphs; Dr. James Kendall for histological prep-
arations of epithelial tissues; Drs. Thomas Clarke
and K. Gopalakrishnan for identification of
stomach contents; Drs. Carl L. Hubbs, Richard
Rosenblatt, John McCosker, Bruce Welton,
Murray Dailey, Scott Johnson, Tyson Roberts,
and John Randall, and Mr. Richard Ellis, for
discussions relating to nomenclature and the sci-
entific importance of the animal; Allan Hart, Re-
becca Brown, and Mary Morioka for scientific
illustrations; Ruth Naftel for organizational sup-

port; and Pam Miike for typing the manuscript.
Particular thanks go to Richard Ellis and John
McCosker for preparation of a preliminary
manuscript which was of great help in the pro-
duction of this final paper.

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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 9, pp. 111-121, 5 figs., 8 tables.

January 17, 1984

THE CRANIAL MORPHOMETRY OF GALAPAGOS TORTOISES

By
Charles R. Crumly1

Department of Zoology and Physiology, Rutgers— The State University,
Newark, New Jersey 07102

ABSTRACT: Saddlebacked tortoises have smaller and slightly broader skulls than non-saddlebacked tortois-
es. Unlike the two types of shells, the skulls of saddlebacked and non-saddlebacked tortoises are difficult to
distinguish, even as large adults. Factor analysis, although suggestive of different growth trends, does not
delineate Geochelone ephippium, G. guntheri, and/or G. nigrita. Discriminant function analysis easily distin-
guishes these species. Geochelone ephippium is discriminated from the other two species on the basis of
overall size and G. nigrita has a smaller exposed basisphenoid than G. guntheri. Species represented by small
samples were compared to Geochelone guntheri, G. ephippium, and G. nigrita by means of a discriminant
function analysis classification procedure. The results suggest that skull variation does not parallel shell
variation.

INTRODUCTION

The classification of Galapagos tortoises (ge-
nus Geochelone) has changed over the years, de-
pending on the prevalent philosophy pertaining
to closely related forms. These insular tortoises
have been considered different species (Van Den-
burgh 1914) or different subspecies (Wermuth
and Mertens 1961, 1977; Crumly 1980, 1982;
MacFarland et al. 1974a, b). Perhaps the best a
priori taxonomic strategy was employed by Fritts
(in press), who considered each geographically
isolated population a separate entity until more
detailed analysis could be completed. These dif-
ferent philosophies, compounded by the fre-
quent lack of accurate locality data, are reflected
in the confusing nomenclatural history of Ga-
lapagos tortoises (Table 1).

1 Present address: Division of Reptiles and Amphibians, De-
partment of Vertebrate Zoology, National Museum of Natural
History, Smithsonian Institution, Washington, D.C. 20560.

Although the nomenclatural status of these
various populations remains changeable, it is clear
that all Galapagos tortoises are more closely re-
lated to each other than to other tortoises. This
interpretation is supported by morphologic anal-
yses (Crumly 1980, 1982; Fritts in press) and
electrophoretic studies (Marlow and Patton
1981).

Despite their close relationships, Galapagos
tortoises exhibit great structural diversity. The
shells best reflect this diversity and are of two
basic types: domed, like those of most other tor-
toise species; and saddlebacked, resembling an
ancient Moroccan saddle. The saddlebacked shell
type seems derived from the domed type, but
Fritts (in press) has noticed subtle differences be-
tween saddlebacked forms that suggest this mor-
phology evolved more than once. Marlow and
Patton (1981) corroborate Fritts's suggestion.
Furthermore, the saddlebacked shell, long con-
sidered unique to certain Galapagos tortoises,
appears independently in Geochelone vosmaeri

[ill]

112

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 9

Pinto A3

San Salvador
fe

°W*2 J\

nzon/^^ San

nta Cruz

San Cristoba
oSanta Fe _/O

ArchipielagodeColon

Espanola

Santa Maria

FIGURE 1 . Present distribution of Geochelone species in the Galapagos Islands. A = abingdonii, B = becki, C = chathamensis,
D = darwini. El = elephantopus, Ep = ephippium, G = guntheri, H = hoodensis, M = microphyes, N = nigrita, Ph = phantas-
ticus, Va = vandenburghi, Vi = vicina, W = wallacei, ? = unnamed form, 1 = domed, 2 = intermediate, 3 = saddlebacked, * =
species probably invalid (from MacFarland et al. 1974a).

of Rodrigues Island in the Indian Ocean (Arnold
1979).

Fritts (in press) showed that island (or volcano)
topography is a reliable predictor of shell shape
in Galapagos tortoises. Based on captive breed-
ing data, he also suggested that shell differences
are genetically determined. The topographic, cli-
matic, and morphologic information combined
in Fritts's model suggests that these tortoises pos-
sess great adaptive plasticity. This plasticity, re-
flected in differences in shell shape, has not been
documented for other anatomical regions.

The purpose of this study is to determine
whether variation in cranial morphometry par-
allels variation in the two shell types, first pro-
posed by Van Denburgh (1914) and corroborated
by Fritts (in press).

MATERIALS AND METHODS

Sixteen measurements (Fig. 2, Table 2) were
recorded from over 100 skulls, listed below. (The
museum acronyms used are those recommended
by Duellman et al. 1978.)

G. abingdonii: CAS 8112; USNM 29269.

G. to?cfc/:CAS8120.

G. chathamensis: CAS 8127, 8128, 8130, 8131,

8133; USNM 29255.
G. darwini: CAS 8106, 8108, 39601.
G. ephippium: AMNH 93383; CAS 8299, 8313,

8358, 8377-8380; MCZ 1 1068; USNM 29309,

29251.
G. guntheri: CAS 8225, 8267, 8413, 8406, 8401,

8399, 8400, 8396, 8415, 8256, 8408, 8405,

8199, 8194, 8210.

CRUMLY: TORTOISE SKULLS

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114

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. Vol. 43. No. 9

FIGURE 2. Measurements taken from Galapagos tortoise skulls (see Table 2 for explanations of abbreviations).

G. hoodensis: CAS 8121, 8122.

G. microphyes: CAS 8158.

G. nigrita: CAS 8381, 8289, 8286, 8385; MVZ
67613-67615, 59528, 67624-67629, 67631-
67633; USNM 104330-104331.

G. phantastica: CAS 8101.

G. vandenburghi: CAS 8141.

G.vicina: CAS 8 179, 8 193, 8 177; USNM 129247.

G. wallacei (probably an invalid form, fide
MacFarland et al. 1974a): CAS 8134.

Geochelone sp. (but definitely Galapagos tortoise
species): AMNH 7288, 42961, 63415, 36420,
36568-36570, 63416; CAS 8298, 8404, 841 1,
8409, 8402, 8377, 8407, 8410, 8403, 8414,
8397, 84 1 2, 8272; Calif. State Univ., Fullerton
Coll. 3 uncat.; FMNH 13523, 1 uncat.; LACM
(Vert. Paleo.) pr 63, pr 58, pr 64; MCZ 46606,
11070, 11069, 32098, 1905, 4668; MVZ
80075; SDSNH 56605, 55458; USNM 65896,
102904, 129393, 15192, 29338, 29305, 29254,
29252, 15190, 15193,29256.

Means and standard deviations were calculat-
ed for each of the 1 6 measurements and corre-
lation coefficients were also calculated.

At the recommendation of Fritts (pers. comm.),
I followed the last thorough taxonomic review

(Van Denburgh 1 9 1 4) in which the different forms
were given species-level designations. The species
names used by Van Denburgh (1914) are fol-
lowed with one exception; G. porteri is consid-
ered a junior synonym of G. nigrita (fide Fritts
in press). Statistical comparisons between island

TABLE 2. SKULL MEASUREMENTS RECORDED FOR GALAP-
AGOS TORTOISES. (All measurements taken with dial calipers
and recorded to nearest 0.01 mm.)

Variable— Description
B— Basicranial length

WAT— Width of skull at anterior tympanic opening
WO— Width between orbits
HN — Height of external narial opening
WN— Width of external narial opening
LB— Length of basisphenoid
WB— Width of basisphenoid
WZ— Width of quadratojugal
WP- Width of postorbital
WS-Width of jugal

DPV — Distance (greatest) from prepalatine foramina (or fo-
ramen, if only one present) to vomer
LP— Length of prootic

WFS — Width of prootic at stapedial foramen
PW— Width of pterygoid waist
APW — Width of anterior premaxillae
PC— Length of sagittal contact of prefrontals

CRUMLY: TORTOISE SKULLS

115

TABLE 3. MEANS AND STANDARD DEVIATIONS FOR 16 VARIABLES IN FIVE GALAPAGOS TORTOISE SPECIES. Measurements are
illustrated in Figure 2 and abbreviations are listed in Table 2. Most sample sizes are small; all measurements are in millimeters.

G. ephippium
(N = 9)

G. guntheri
(N= 15)

G. nigrita
(N=18)

G. vicina
(N = 4)

G. chathamensis
(N = 6)

Variable

96.7

11.4

128.0

21.4

121.5

39.2

109.0

49.2

98.1

27.3

WAT

73.9

9.2

106.6

19.9

98.4

31.8

86.0

38.9

80.4

25.0

25.1

3.1

35.4

7.3

37.0

13.3

28.4

12.6

28.4

7.7

12.5

2.0

18.6

3.4

18.6

6.0

16.1

7.3

13.9

4.1

17.0

2.2

25.1

4.4

23.1

7.7

21.3

9.2

18.5

4.7

13.3

3.2

18.7

4.2

14.7

4.3

18.1

8.9

14.7

5.6

14.6

2.2

19.1

4.3

17.1

4.6

15.8

7.2

13.8

3.4

9.3

3.6

14.3

4.4

13.3

4.7

12.6

6.8

10.1

3.5

7.0

2.2

9.5

2.5

9.0

3.5

8.8

4.5

7.3

3.0

7.3

2.3

12.0

3.0

9.5

4.0

9.6

5.1

7.9

3.1

DPV

3.2

0.8

4.2

0.8

4.2

1.5

3.7

2.4

3.1

1.0

14.1

2.1

21.0

5.6

18.1

6.5

14.8

6.6

15.2

3.7

WFS

10.0

2.0

16.0

6.1

12.8

6.1

8.9

6.3

12.5

6.2

19.2

1.8

25.9

5.1

26.1

8.4

21.9

8.6

19.0

4.0

APW

10.5

1.8

15.2

2.4

14.1

4.9

11.8

8.0

10.5

3.0

8.6

1.5

10.6

4.0

13.3

5.4

8.5

3.9

8.2

3.9

populations were hampered by incomplete lo-
cality data; 50 of 1 1 6 specimens (43%) examined
possessed doubtful or unknown locality data. The
specimens without locality data were readily
identified as Galapagos tortoises, but could not
be identified to species without locality data.
These specimens were used in the computation
of correlation coefficients and in factor analysis,
but could not be used in other statistical proce-
dures.

To facilitate my analyses, populations were
combined based on the shell types advocated by
Van Denburgh (1914) and Fritts (in press). Thus,
the saddlebacked forms (G. abingdonii [N = 2],
G. phantastica [N = 1], G. becki [N = 1], G.
hoodensis [N = 2], and G. ephippium [N = 9])
were combined, yielding a sample of 1 5 individ-
uals. The non-saddlebacked forms (intermediate
and domed shells of Van Denburgh 1914) were
also combined, forming a larger sample of 48
individuals (G. chathamensis [N = 6], G. dar-
wini [N = 2], G. guntheri [N = 15], G. micro-
phyes [N = 1], G. nigrita [N = 18], G. vicina
[N = 4], and G. vandenburghi [N = 1]). These
larger samples were then compared to determine
whether cranial variation mirrored the already
well known shell variation. Comparisons were
also made among G. ephippium, G. guntheri, and
G. nigrita to determine whether noncombined
and combined samples contained the same mag-
nitude of variation.

The Statistical Package for the Social Sciences
(SPSS) was used on the WYLBUR facility at the
Campus Computer Information Service (CCIS)
at Rutgers— The State University for initial data
examination. Final statistical analyses were ac-
complished using SPSS programs available

TABLE 4. MEANS AND STANDARD DEVIATIONS FOR SADDLE-
BACKED TORTOISES REPRESENTED BY SPECIMENS OF FIVE SPECIES
AND NON-SADDLEBACKED TORTOISES REPRESENTED BY SPECI-
MENS OF SEVEN SPECIES. Measurements are illustrated in Figure
2 and abbreviations are listed in Table 2; all measurements
are in millimeters.

Saddlebacked
(N= 15)

Nonsaddlebacked
(N = 48)

Variables

98.9

15.9

116.5

37.1

WAT

75.8

13.0

96.0

30.4

26.4

4.6

32.9

11.2

13.2

2.9

17.1

5.6

17.5

2.4

22.6

7.2

13.8

3.2

16.0

5.7

14.5

2.0

17.1

5.5

9.5

3.4

13.1

5.3

7.1

2.2

8.8

3.5

7.3

2.3

9.9

3.9

DPV

2.9

1.2

3.9

1.4

14.4

2.7

18.4

6.2

WFS

10.1

1.8

13.7

6.4

20.1

2.9

24.6

7.8

APW

10.5

1.7

13.2

4.5

7.9

1.4

10.2

5.2

116

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 9

TABLE 5. CORRELATION COEFFICIENTS BETWEEN ALL THE SKULL MEASUREMENTS ILLUSTRATED IN FIGURE 2 AND ABBREVIATED
IN TABLE 2. All specimens measured are combined into a single sample. Nevertheless, all coefficients are significant to at least
the P = 0.05 level.

Variable

WAT

DPV

LP WFS PW APW

WAT

.98

.94

.96

.94

.93

.97

.96

.93

.95

.88

.86

.79

.84

.88

.92

.87

.88

.92

.87

.84

.86

.81

.87

.78

.84

.87

.86

.88

.83

.87

.79

.80

.88

.91

.92

.87

.88

.91

.86

.89

.84

DPV

.69

.64

.72

.60

.68

.64

.66

.95

.94

.91

.94

.85

.91

.86

.84

.91

.63

WFS

.85

.78

.77

.87

.90

.91

.78

.88

.85

.83

.69

90 82

APW

.93

.89

.91

.94

.84

.89

.82

.83

.89

.68

.86 .82 85

.49

55 50 54 59

through the Office of Computer Services (OCS)
at the Smithsonian Institution. Simple descrip-
tive statistics, linear regression, factor analysis,
and stepwise discriminant analyses were used to
summarize observed cranial variation.

RESULTS

Geochelone ephippium appears to have the
smallest skull and G. guntheri the largest skull
of Galapagos tortoises (Table 3), but when max-
imum basicranial lengths (mean plus two stan-
dard deviations) are compared, G. nigrita ap-
pears to possess the largest skull (Bmax = 171 mm
for G. guntheri, 200 mm for G. nigrita). The
efficacy of this procedure is in some doubt since
the Bmax for G. vicina exceeds that of G. nigrita,
even though no skull of the former is anywhere
near as large as the latter. This may be the prod-
uct of a small sample size for G. vicina, repre-
sented by only four specimens. The largest skulls
in these samples are G. ephippium, 1 14.0 mm;
G. guntheri, 157.7 mm; G. nigrita, 157.6 mm;
and G. vicina, 142.7 mm. The G. nigrita sample
includes the two smallest tortoises measured,
which depresses the mean basicranial length and
elevates the standard deviation.

Combined samples clearly show a size differ-
ential between saddlebacked and domed tortois-
es; saddlebacked tortoises have smaller skulls.
This is supported by all 16 variables (see Table
4).

All correlation coefficients were significant to
at least the P = 0.05 level (Table 5). Some vari-

ables, however, did not correlate as highly with
other variables. Examples include PC, DPV, and
LB. Because intervariable correlation was so high,
linear regression showed slight, if any, tendency
toward curvilinearity. The intercepts for saddle-
backed forms were lower than the intercepts for
non-saddlebacked forms, reflecting the differ-
ence in size between the two groups. Slopes, how-
ever, were practically identical. As an example,
linear equations relating WO to LB for saddle-
backed and non-saddlebacked tortoises have
slopes of 1.38 and 1.37, respectively, whereas
intercepts are 7.92 and 1 1.74, respectively (r =
0.75 for saddlebacks and 0.69 for nonsaddle-
backs, P < 0.005 for both).

Factor analysis yielded three factors, the first
accounted for almost 95% of the data variance
(see Table 6). Before rotation all 16 variables
correlated most highly with this first factor. Ro-
tation simplifies vectors derived by the analysis
procedure and is necessary because factor anal-
ysis problems have more than one solution. There
are two general rotation techniques: orthogonal
and oblique. Orthogonal rotation solutions de-
rive vectors along axes of data variation that are
perpendicular to one another and thus uncorre-
lated. Oblique techniques, on the other hand, do
not require that vectors be orthogonal, so vectors
can be correlated. Even after varimax rotation,
an orthogonal technique that simplifies the col-
umns of a factor matrix by maximizing factor-
variable loadings, 12 of the 16 variables correlate
most highly with factor one. Varimax rotation

CRUMLY: TORTOISE SKULLS

117

C© 00,

Factor 2

-2-

Factor 3

FIGURE 3. A plot of factor scores for factors two and three. Geochelone nigrita (solid circles), G. guntheri (cross-hatched
circles) and G. ephippium (open circles). When factor scores for all tortoises are plotted there is a prominent trend from the
lower-left to upper-right quadrant. Although this general trend for all tortoises is suggestive of a positive trend toward increased
snout elongation with increased robustness (as illustrated by G. nigrita), the points for G. ephippium and G. guntheri show a
negative relationship between robustness and snout elongation.

was chosen because it maximizes the variation
accounted for by the factor vectors without all
the variables loading highly on the same factor,
as occurs in quartimax rotation.

Identifying vectors of data variation is spec-
ulative; but it seems likely that factor one sum-
marizes variation in size. Thus, 95% of the vari-
ation in Galapagos tortoise skulls may be the
result of variation in size. The other two factors
are more difficult to interpret, partly because so
little variation (only 5%) is summarized by these
factors. Factor two summarizes variation in cra-

TABLE 6. STATISTICS PRODUCED BY FACTOR ANALYSIS USING
VARIMAX ROTATION. All specimens were included in this anal-
ysis. Abbreviations used in the summarized factor matrix are
listed in Table 2. Eigenvalues are measures of the relative
importance of the factors.

Factor

Eigenvalue

13.19

0.47

0.28

% Variation

94.6

3.4

2.0

Summarization of

WFS 0.82

WP 0.63

PC 0.75

Factor Matrix

LB 0.80

DPV 0.60

HN 0.51

LP 0.78

PW 0.53

WAT 0.76

WO 0.53

WS 0.76

WZ 0.72

B 0.76

WB 0.75

WN 0.73

HN 0.72

nial width and the width of skull arches, em-
phasizing WO, WP, WZ, PW, and DPV. There-
fore, factor two could be identified as some
measure of robustness. Factor three, emphasiz-
ing PC and HN, suggests there is variation in the
anterior part of the skull. A high factor three
score results from an increase in PC and HN.
This results from elongating the anteromedial
portion of the triturating surface, which concom-
itantly yields a longer skull.

A bivariate plot of the second and third factor
scores for G. nigrita, G. guntheri, and G. ephip-
pium (Fig. 3) indicates that as skulls become more
robust, the anterior nasal part of the skull elon-
gates; as robustness increases the skull becomes
relatively longer. However, examining the indi-
vidual points for G. guntheri and G. ephippium
suggests just the opposite; as robustness increases
elongation decreases. This negative relationship
seems more pronounced in G. guntheri.

Three separate discriminant function analyses
were done: one for G. nigrita, G. ephippium, and
G. guntheri; one for the combined samples; and
one comparing small samples to larger samples.
In the first analysis, the three forms were distin-
guished by two factors (Table 7). Factor one sum-
marized variation in 14 of the 16 variables but
accounted for only 54.9% of the data variance.
A high canonical correlation coefficient and a low
Wilks's lambda indicate that this factor is good

118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. Vol. 43, No. 9

Jill

-6

-2

2ee 4

9 99 _2

9
a9

FIGURE 4. A plot of the discriminant scores derived from
an analysis that included Geochelone guntheri, G. nigrita, and
G. ephippium. Discriminant factor one is the horizontal axis
and discriminant factor two is the vertical axis. Statistical sep-
aration of these three populations is marked. High positive
scores along the horizontal axis indicate small size, whereas
high positive scores along the vertical axis indicate a poorly
exposed basisphenoid. Upper case letters indicate group cen-
teroids.

at distinguishing groups. Geochelone ephippium
is separated from the other two forms by this
factor. The discriminating variables are nega-
tively correlated to factor one, and the species
with a small skull is differentiated from the two
species with large skulls, suggesting that factor

TABLE 7. STATISTICS PRODUCED BY A DISCRIMINANT
ANALYSIS OF G. ephippium, G. guntheri AND G. nigrita. Ab-
breviations are listed in Table 2. Eigenvalues are measures of
the relative importance of the factors; high canonical corre-
lation coefficients (near 1) and low Wilks's lambdas (near 0)
indicate that factors are good discriminators.

Discriminant
function

Eigenvalue

7.06

5.82

% Variation

54.8

45.2

Canonical correlation

0.94

0.92

Wilks's lambda

0.02 (P = 0.003)

0.15(/>=0.02)

Pooled within groups

HN -0.22

LB -0.21

correlations bet.

LP -0.20

WB -0.17

canonical

WN -0.19

discr. fncts. &

WAT-0.19

discr. variables

APW-0.18

WO -0.18

Groups delineated

G. ephippium

G. nigrita

from others

from G.

guntheri

FIGURE 5. A histogram illustrating the results of a discrim-
inant analysis of the saddlebacked and non-saddlebacked forms.
The saddlebacked species are in the upper histogram, the non-
saddlebacked species are in the lower histogram. The arrows
indicate the median in each class. The discriminant scores (high
positive scores indicate large size) are on the lower axis and
the number of individuals are represented by left-hand axis.
Although the saddlebacked and non-saddlebacked forms are
clearly different sizes, there is significant overlap.

one is an inverse measure of size. Factor two,
which also has a high canonical correlation coef-
ficient and a low Wilks's lambda, distinguishes
G. nigrita from G. guntheri and accounts for the
remaining variation in the data. Two variables
are highly correlated with this second factor, LB
and WB. Geochelone nigrita has high positive
values for discriminating factor two, indicating
that the basisphenoid is poorly exposed. Figure
4 graphically illustrates the completeness of sep-
aration.

Standardized canonical discriminant function
coefficients are available upon request. These
coefficients can be used to calculate discriminant
scores for individual specimens whose identity
is unknown; but choices are restricted to the pop-
ulations originally compared (in this case G.
ephippium, G. guntheri, or G. nigrita).

The second discriminant analysis applied to
the combined samples. Because only two groups
were analyzed, a single discriminating factor was
computed. The Wilks's lambda was not low, sug-
gesting that the two groups cannot be easily dis-
tinguished. The size differential between saddle-
backed and non-saddlebacked tortoises is readily
apparent (Fig. 5). Standardized canonical dis-
criminant function coefficients are available upon
request.

The third discriminant analysis compared

CRUMLY: TORTOISE SKULLS

TABLE 8. CLASSIFICATION RESULTS OF A DISCRIMINANT ANALYSIS CLASSIFICATION PROCEDURE. Individual specimens were
classified to one of three species: G. ephippium (a saddlebacked species), G. guntheri (an intermediate form) or G. nigrita (a
domed form). Asterisk indicates invalid taxon (fide MacFarland et al. 1974a).

Shell type

Trivial name

Sex

Mus. no.

Classified as

domed

vandenburghi

CAS 8141

ephippium

intermediate

chathamensis

CAS 8 133

ephippium

intermediate

chathamensis

CAS 8131

ephippium

intermediate

chathamensis

USNM 29255

ephippium

intermediate

chathamensis

male

CAS 8 127

ephippium

intermediate

chathamensis

CAS 8 130

ephippium

intermediate

chathamensis

CAS 8 128

ephippium

intermediate

darwini

female

CAS 8 106

ephippium

intermediate

darwini

male

CAS 8 108

guntheri

intermediate

microphyes

male

CAS 8 158

guntheri

intermediate

vicina

male

CAS 8 179

ephippium

intermediate

vicina

female

CAS 8 193

ephippium

intermediate

vicina

USNM 129247

ephippium

intermediate

vicina

male

CAS 8 177

guntheri

intermediate

wallacei*

male

CAS 8 134

guntheri

saddlebacked

abingdonii

USNM 29269

guntheri

saddlebacked

abingdonii

male

CAS 81 12

guntheri

saddlebacked

becki

female

CAS 8 120

ephippium

saddlebacked

hoodensis

male

CAS 8121

ephippium

saddlebacked

hoodensis

female

CAS 8 122

ephippium

saddlebacked

phantastica

male

CAS 8101

guntheri

small samples of tortoise species to large sam-
ples. Small samples were classified by the dis-
criminant function classification procedure to one
of three species (G. guntheri, G. ephippium, G.
nigrita). The results of this procedure are sum-
marized in Table 8. Some species with inter-
mediate shell types (fide VanDenburgh 1914) were
classified as saddlebacked species (e.g., G. chath-
amensis was classified as G. ephippium), whereas
other species with intermediate shell types were
classified as G. guntheri, an intermediate form.
No species was classified as a domed form. Skull
variation did not parallel shell variation in any
meaningful way.

DISCUSSION

Small sample sizes and the paucity of accurate
locality data limit the utility of this study. There-
fore, samples were combined. (Thorpe, 1976,
discusses the ramifications of such procedures.)
Because most of the specimens in the United
States were examined, this limitation cannot be
overcome without costly and time-consuming
removal of skulls from skins and stuffed speci-
mens of known provenance.

The choice of a putative ancestral morphotype
makes an enormous difference in how one in-
terprets evolutionary processes, patterns, and

mechanisms. The size of the ancestral Galapagos
tortoise is not known. Auffenberg (1971) be-
lieved that the fossil Geochelone hesterna was a
likely ancestral candidate for Galapagos tortoises
as well as Geochelone chilensis from Argentina.
The skull of G. hesterna is very much like a
Galapagos tortoise skull. Although it is not as
large as that of the largest of Galapagos domed
tortoises, it is larger than that of the small sad-
dlebacked tortoises. Thus, I favor an interme-
diate-sized ancestor for Galapagos tortoises, per-
haps something smaller than G. guntheri. If so,
then G. nigrita is the result of continued gigan-
tism and G. ephippium is the result of dwarfism.

Why is there such flimsy coincidence between
shell variation and cranial variation in Galapa-
gos tortoises? Zangerl and Johnson (1957) and
Zangerl (1969) have intimated that much of the
shell variation observed in most species has little
effect on an individual's survival or fitness. Fritts
(in press) has shown the contrary for Galapagos
tortoises. But this selection on shell morphology
does not seem to apply to skull morphology. What
other selective factors could be molding skull
morphology?

I tend to agree with Bramble (1971), who felt
that biomechanical constraints on chewing are
the primary sources of selection upon turtle skulls.

120

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43. No. 9

How strong are these selective forces? Selective
factors imposed by diet are known to be espe-
cially strong in other animals from Galapagos.
For example, Boag and Grant (1981) discovered
that finches in Galapagos experienced intense se-
lection upon beak size and shape as the result of
a one-year drought. Because of the long life of
tortoises and their ability to survive long periods
without food or water, however, short-term en-
vironmental changes such as those described by
Boag and Grant are unlikely to affect tortoises
as severely.

Another source of selective pressure is possi-
ble. During intraspecific agonistic encounters
(Fritts, pers. comm.), the victor is the individual
capable of raising its head the highest. Could
apparent head width also affect the outcome of
these battles? Interestingly, the relative head
width of Geochelone guntheri and G. ephippium
increases with size. These tortoises inhabit low
dry islands (or parts of islands) where carrying
capacities of the habitat may be lower and in-
traspecific competition therefore higher. In con-
trast, relative head width in G. nigrita decreases
with size. This tortoise lives on a higher moist
island where carrying capacities may be higher
and intraspecific competition may not be as in-
tense. Also, this apparent decrease in relative
width actually accompanies an increase in the
length of the masticatory surface area, perhaps
allowing more efficient mastication.

ACKNOWLEDGMENTS

I thank L. Barnes (LACM), R. Crombie
(USNM), R. Drewes (CAS), G. Foley (AMNH),
T. Fritts (SDSNH), A. Leviton (CAS), H. Marx
(FMNH), W. Presch, A. Resetar (FMNH), J. Ro-
sado (MCZ), H. Voris (FMNH), D. Wake (MVZ),
E. Williams (MCZ), G. Zug (USNM), and R.
Zweifel (AMNH) for making specimens avail-
able. I also thank R. Crombie, C. Ernst, T. Fritts,
S. McDowell, E. Meyer, K. Miyata, D. Stead-
man, and G. Zug for reading parts or all of the
manuscript and providing helpful suggestions.
Funding, for which I am most thankful, came
from a Sigma Xi Grant-In-Aid of Research, the
Alvarado Community Hospital Research Foun-
dation, the Vertebrate Zoology Reserve Fund of
San Diego State University, and the Theodore
Roosevelt Memorial Fund of the American Mu-
seum of Natural History. Data analysis, begun
at Rutgers— The State University with Depart-

ment of Zoology and Physiology support, was
completed with the assistance of the Office of
Computer Services during my tenure as a Smith-
sonian Predoctoral Fellow at the National Mu-
seum of Natural History.

LITERATURE CITED

ARNOLD. E. N. 1979. Indian Ocean giant tortoises: their sys-
tematics and island adaptations. Phil. Trans. R. Soc. Lond.
(8)286:127-45.

AUFFENBERG, W. 1971. A new fossil tortoise, with remarks
on the origin of South American testudinines. Copeia 1971:
106-17.

BOAG, P. T., AND P. R. GRANT. 1981. Intense natural selec-
tion in a population of Darwin's finches (Geospizinae) in the
Galapagos. Science 214:82-85.

BRAMBLE, D. M. 1971. Functional morphology, evolution,
and paleoecology of gopher tortoises. Ph.D. thesis, Univ.
Calif., Berkeley. 341 p.

CRUMLY, C. R. 1980. The cranial osteology and evolution
of the tortoise genus Geochelone (Testudines, Testudinidae).
M.S. thesis, San Diego State Univ. 338 p.

. 1982. A cladistic analysis of Geochelone using cranial

osteology. J. Herpetol. 16(3):2 15-34.

DUELLMAN, W. E., T. FRITTS, AND A. LEVITON. 1978. Mu-
seum acronyms. Herp. Rev. 9(l):5-9.

FRITTS, T. H. [in press]. Morphometrics of Galapagos tor-
toises: evolutionary implications.

GARMAN, S. 1917. The Galapagos tortoises. Mem. Mus.
Comp. Zool. 30:262-96.

GUNTHER, A. 1875. Description of the living and extinct
races of gigantic tortoises. Parts I and II: The tortoises of
the Galapagos Islands. Phil. Trans. R. Soc. Lond. (B) 165:
251-84.

. 1877. The gigantic land tortoises (living and extinct)

in the collection of the British Museum. British Museum.
London. 96 p.

MACFARLAND, C. G., J. VILLA, AND B. TORO. 1974a. The
Galapagos giant tortoises (Geochelone elephantopus). Part I:
The status of the surviving populations. Biol. Conserv. 6:
118-33.

. 1974b. The Galapagos giant tortoises (Geochelone

elephantopus). Part II: Conservation methods. Biol. Con-
serv. 6:198-212.

MARLOW, R., AND J. L. PATTON. 1981. Biochemical rela-
tionships of the Galapagos tortoises (Geochelone elephan-
topus). J. Zool., London 195:413-22.

PRITCHARD, P. C. H. 1979. Encyclopedia of turtles. T. F. H.
Publishers. 895 p.

ROTHSCHILD, L. 1901. A new land tortoise from the Gala-
pagos Islands. Nov. Zool. 8:372.

. 1902. Description of a new species of gigantic land

turtle from the Galapagos Islands. Nov. Zool. 9:619.

. 1903. Description of a new species of gigantic tortoise

from Indefatigable Island. Nov. Zool. 10:119.

. 1915. The gigantic land tortoises of the Galapagos

Islands in the Tring Museum. Nov. Zool. 22:403-17.

THORPE, R. S. 1976. Biometrical analysis of geographic vari-
ation and racial affinities. Biol. Rev. 51:407-52.

VAN DENBURGH, J. 1907. Expedition of the California Acad-
emy of Sciences to the Galapagos Islands, 1905-1906. Part

CRUMLY: TORTOISE SKULLS 1 2 1

I: Preliminary descriptions of four new races of gigantic land — . 1977. Liste der rezenten Amphibien un Reptilien.

tortoises from the Galapagos Islands. Proc. Calif. Acad. Sci. Testudines, Crocodylia, Rhyncocephalia. Das Tierrich 100:

1:1-16. 1-174.

1914. Expedition of the California Academy of Sci- ZANGERL, R. 1969. The turtle shell. Pages 31 1-339 in Gans,

ences to the Galapagos Islands, 1905-1906. Part X: The C, A. d'A. Bellaris, and T. S. Parsons. Biology of the Rep-
gigantic land tortoises of the Galapagos Archipelago. Proc. tilia, vol. 1 , Academic Press, New York.
Calif. Acad. Sci., ser. 4, 2:203-374. — , AND R. G. JOHNSON. 1957. The nature of shield
WERMUTH, H., AND R. MERTENS. 1961. Schildkroten, kro- abnormalities in the turtle shell. Fieldiana: Geol. 10:341-
kodile, und bruckenechsen. Gustav Fischer Verlag, Jena. 62.
422 p.

CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, California 941 18

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 10, pp. 123-140, 8 figs., 1 table

January 17, 1984

THE STATUS OF TRYPOXYLON FIGULUS

(LINNAEUS, 1758), MEDIUM DE BEAUMONT,

1945, AND MINUS DE BEAUMONT, 1945

(HYMENOPTERA: SPHECIDAE)

By
Wojciech J. Pulawski

California Academy of Sciences, Golden Gate Park, San Francisco, California 94118

ABSTRACT: Trypoxylon figulus (Linnaeus, 1758), medium de Beaumont, 1945, and minus de Beaumont,
1945, currently confused under the name figulus, are separated on the basis of newly discovered characters.
/. majus Kohl, 1883, figulus barbarum de Beaumont, 1957, and figulus yezo Tsuneki, 1956, are newly
synonymized with figulus, and figulus koma Tsuneki, 1956 is newly synonymized with minus. Neotypes are
designated for Sphex fuliginosus Scopoli, 1763, and Trypoxylon majus Kohl, 1883, both synonyms of figulus,
and a lectotype is designated for Trypoxylon rubi Wolf, 1959, a synonym of medium.

INTRODUCTION

De Beaumont (1945) was first to observe that
Trypoxylon figulus of European authors actually
consisted of three phena. Their status has been
controversial over the years. De Beaumont ( 1 945,
1964a) and Richards (1980) called them vari-
eties, Bliithgen (1951) gave them species rank,
and Wolf (1959) and Bohart and Menke (1976)
treated them as subspecies. The last interpreta-
tion is untenable, since the three phena are large-
ly sympatric. Tsuneki (1981) regarded medium
as a good species, characterized by both external
and genitalic characters, and considered minus
as a simple form of figulus.

According to Valkeila (1961), specimens reared
from one nest mostly are one phenon, but he
reported that two phena (e.g., majus and minus)
are found in some nests. He concluded that all
three are individual variants of one species. Un-
fortunately, Valkeila's data cannot be verified. I
have examined all of his specimens, which are

presently kept at Helsinki University. Some
specimens have identification labels by de Beau-
mont, but not a single label refers to nests or cells
from which specimens were reared. Possibly
Valkeila misidentified some specimens, but this
cannot be determined because his identification
labels give the name figulus only, without ref-
erence to form or varietal names. Another pos-
sible explanation is that offspring of two nests
were accidentally confused.

A thorough examination of the three phena,
based on more than 3800 specimens from many
countries, convinced me that actually they are
good species. My opinion is based on the follow-
ing evidence:

1. Morphology. Although some males of fi-
gulus and minus cannot be distinguished with
certainty, females are separated by structural gaps
and do not intergrade; also the male of medium
is easily recognized by its peculiar gonoforceps.
Some previously unnoticed characters (antero-

[123]

124

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 10

ventral mesothoracic process, length of thoracic
vestiture, female hindcoxal pit) are especially
helpful in recognizing the three species.

2. Rearing. Of 9 1 specimens reared from trap
nests in Liege, Belgium, by A. Jacob-Remacle,
and examined by me, all are minus (7 2, 5 3 reared
in 1976, 44 2, 35 $ reared in 1981). This result
contradicts Valkeila's conclusions: If the three
phena really are variants of one species, then one
would expect some variation of phena in the
Liege trap nest material.

3. Geographic distribution. All three species
are largely sympatric, but only figulus and me-
dium have been found in Great Britain, only
figulus and minus in the Iberian Peninsula, and
only figulus in North Africa and North America.
If the three phena were just individual forms of
one species, some variation should have oc-
curred in North American populations, and all
three phena should have been observed in Great
Britain (all three do occur in the Netherlands,
where climatic and ecological conditions are
practically identical to those in England).

4. Habitat preference. Of 429 specimens col-
lected by J. Leclercq in waterbowl traps in Liege,
Belgium, in 1980 and 1981 (and examined by
me), 4 are medium and the remainder are minus
(202 2, 223 <5). Not a single figulus was trapped,
in spite of the fact that this species is common
in Belgium.

I fully agree with Tsuneki (198 1) that some of
the previously used characters are not reliable.
For example, the mesopleural punctures of fi-
gulus are dense and well denned according to de
Beaumont (1945, 1964a), but in the smallest
males they actually are as sparse and minute as
in most minus. Such characters have not been
used here.

Proper mounting is critical for studying the
species considered. For example, the anteroven-
tral mesothoracic region must not be damaged
by the pin. When pinning the specimens, one
should insert the pin so that it passes through
the membrane between the mesothorax and fore-
coxae (which then extend laterad). In this posi-
tion, the anteroventral mesothoracic region is
easily visible. Unfortunately, many European
collectors mount their specimens on cardboard
rectangles with glue or minutiae (venter down
rather than on a side), and such specimens must
be relaxed and remounted before examination.

In the text below the locality records are ar-

ranged according to current administrative di-
visions for each country except Sweden and Fin-
land, for which biogeographic provinces have
been used. Localities given on specimen labels
but not found on available maps or in gazeteers
have not been considered.

An exclamation mark preceding the word Ho-
lotype or Neotype in the bibliographic citations
indicates that the type has been examined.

SOURCES OF MATERIAL

The specimens examined came from institu-
tional and private collections listed below. The
acronyms preceding the names are the abbrevi-
ations by which these collections are referred to
in the text.

AKM: Aimo K. Merisuo, Turku, Finland

AWE: Father Andreas W. Ebmer, Linz, Austria

BB: Padre Bruno Bonelli, Cavalese, Italy

BMNH: British Museum (Natural History), London, England
(Mr. C. R. Vardy)

CAS: California Academy of Sciences, San Francisco, Cali-
fornia (W. J. Pulawski)

CNC: Canada National Collection of Insects, Arachnids and
Nematods, Biosystematics Research Institute, Ottawa, On-
tario

CU: Cornell University, Department of Entomology and Lim-
nology, Ithaca, New York (Dr. L. L. Pechuman)

DBB: Major Donald B. Baker, Ewell, Surrey, England

DEI: Institut fur Pflanzenschutzforschung der Akademie der
Landwirtschaftswissenschaften der DDR, Zweigstelle Ebers-
walde, Abteilung Taxonomie der Insekten (formerly
Deutsches Entomologisches Institut), Eberswalde-Finow (Dr.
J. Oehlke)

FIS: Forschungsinstitut Senckenberg, Frankfurt am Main, Fed-
eral Republic of Germany (Dr. J.-P. Kopelke)

FJS: Seftor Francisco J. Suarez, Almeria, Spain

FSAG: Faculte de Sciences Agronomiques, Gembloux, Bel-
gium (Dr. J. Leclercq)

GP: Signer Guido Pagliano, Turin, Italy

GVR: Mr. Gerard van Rossem, Wageningen, The Netherlands

HD: Dr. Holger Dathe, Forschungsstelle fur Wirbeltierfor-
schung, Berlin, German Democratic Republic

HW: Herr Heinrich Wolf, Plettenberg, Federal Republic of
Germany

HY: Helsingin Yliopisto (=University of Helsinki), Depart-
ment of Agricultural and Forest Zoology, Finland, including
E. Valkeila collection (Dr. Martti Koponen)

IEE: Institute Espanol de Entomologia, Madrid, Spain (Dr. E.
Mingo Perez)

JG: Dr. Joseph Gusenleitner, Linz, Austria

KMG: Mr. Kenneth M. Guichard, % British Museum (Natural
History), London, England

K.S: Professor Dr. Konrad Schmidt, Zoologisches Institut der
Universitat, Karlsruhe, Federal Republic of Germany

KT: Professor Katsui Tsuneki, Mishima, Japan

LEM: Lyman Entomological Museum & Research Laboratory,
Ste. Anne de Bellevue, Quebec, Canada (Dr. A. Finnamore)

MGA: Muzeul de Istorie Naturala Grigore Antipa, Bucharest,
Rumania (Mrs. X. Scobiola Palade)

PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS

125

TABLE 1. ANCESTRAL AND DERIVED CHARACTER STATES OF THREE SPECIES IN THE GENUS TRYPOXYLON.

Character

Ancestral

Derived

1 . Thoracic pilosity
2. Antero ventral mesothoracic process
3. Free margin of female clypeus
4. Female hindcoxal pit
5. Sete of hindcoxal pit
6. Male apical flagellomere
7. Gonoforceps process

shorter (as in medium)
absent
straight or sinuate
circular
evenly distributed
shorter (as in medium)
absent

longer (as in figulus)
present
concave
oblong
channel-like structure
longer (as in figulus)
present

MHNG: Museum d'Histoire Naturelle de Geneve, Switzerland
(Dr. Cl. Besuchet)

MSNM: Museo Civico di Storia Naturale, Milano, Italy (Dr.
C. Leonardi)

MCZ: Museum of Comparative Zoology, Harvard University,
Cambridge, Massachusetts

NHMB: Naturhistorisches Museum Bern, Bern, Switzerland
(Dr. H. D. Volkart)

NHMV: Naturhistorisches Museum, Vienna, Austria (Dr. M.
Fischer)

NRS: Naturhistoriska Riksmuseet, Stockholm V, Sweden (Mr.
S. Erlandsson)

RMNH: Rijksmuseum van Natuurlijke Historic, Leiden, The
Netherlands, including collections of J. P. van Lith and P.
M. F. Verhoeff (Dr. K. van Achterberg)

SFG: Dr. Severiano Fernandez Gayubo, Departamento de
Zoologia, Universidad de Salamanca, Spain

SMT: Staatliches Museum fur Tierkunde, Dresden, German
Democratic Republic (Dr. Regine Eck)

TMB: Termeszettudomanyi Muzeum, Budapest, Hungary (Dr.
J. Papp)

TN: Mr. Toshiaku Nambu, Yorii-machi, Saitama Prefecture,
Japan

USNM: United States National Museum (Smithsonian Insti-
tution), Washington, D.C.

VH: Dr. Volk Haeseler, Universitat Oldenburg, Oldenburg,
Federal Republic of Germany

VLK: Dr. Vladimir L. Kazenas, Zoological Institute, Kazakh
Academy of Sciences, Alma Ata, USSR

WJP: Wojciech J. Pulawski, San Francisco, California

WSU: Washington State University, Department of Ento-
mology, Pullman, Washington

ZMB: Museum fur Naturkunde an der Humboldt Universitat
zu Berlin, German Democratic Republic (Dr. F. Koch)

ZMH: Zoologisches Institut und Zoologisches Museum der
Universitat Hamburg, Federal Republic of Germany (Dr.
R. Abraham)

ZMK: Zoological Museum, Copenhagen, Denmark (Dr. O.
Lomholdt)

ZMMU: Zoological Museum, Moscow State University, Mos-
cow, USSR (Dr. L. V. Zimina, via Dr. A. P. Rasnitsyn)

ZMUB: Zoological Museum, University of Bergen, Norway
(Dr. Lita Greve Jensen)

ZSM: Zoologische Staatssammlung Milnchen, Federal Repub-
lic of Germany (Dr. E. Diller)

ACKNOWLEDGMENTS

Study of the Linnean type of Sphex figulus at
the Burlington House, London, was possible ow-

ing to the kind assistance of Michael C. Day of
the British Museum (Natural History). North
American specimens of figulus belonging to var-
ious U.S. and Canadian institutions listed above
(except for LEM) were kindly forwarded by Rolin
C. Coville, University of California, Berkeley,
who had them on loan. Arnold S. Menke and
Eric E. Grissell critically reviewed the manu-
script and made many valuable suggestions. Da-
vid H. Kavanaugh commented on the phyloge-
netic trees. Mary Ann Tenorio drew the
phylogenetic schemes and distributional maps,
and Donald J. Becker took the photographs with
a Hitachi S-520 scanning electron microscope.

PHYLOGENETIC RELATIONSHIPS

Reconstructing phyletic relationships between
three isolated species of a large genus like Try-
poxylon is precarious, because polarities of mor-
phological transformations can easily be misin-
terpreted. With this restriction in mind, I
nevertheless think it worthwhile to analyze the
relationships between figulus, medium, and mi-
nus. Their ancestral and derived character states,
based on outgroup comparisons, are shown in
Table 1.

Based on the above table, the three possible
phylogenetic trees (only dichotomic trees are
considered) are as shown in Fig. 1 . Tree B is the
most probable, since no single derived character
state is shared by any two of the three species in
the schemes A and C. Furthermore, trees A and
C imply a parallel development of the elongate
male flagellomere XI in figulus and minus, an
unlikely event.

KEY TO THE SPECIES

1 . Female: clypeal free margin evenly concave
between orbit and median projection (Fig.
7A); mesopleural setae around scrobe
shorter than midocellar diameter; hindcox-

126

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 10

FIGURE 1 . Hypothetical phylogenetic trees of Try poxy Ion figulus (fig), medium (med), et minus (min). Open circles: generalized
character states. Solid circles: derived character states. Numbers refer to characters listed on p. 125.

al pit nearly circular, its setae forming pe-
culiar, channel-like structure (Fig. 7B-D).
Male: maximum length of flagellomere X
0.75-0.9 times its width; maximum length
of flagellomere XI generally 2.0-2.2 times
its basal diameter (occasionally 2.4 times);
gonoforceps with externoventral expansion

at about midlength (Fig. 7E, F)

medium de Beaumont

- Female: clypeal free margin (Fig. 2A, 5A)
sinuate to nearly straight, except concave
in occasional western European figulus in
which mesopleural setae around scrobe are
generally longer than midocellar diameter,
and hindcoxal pit is generally oblong; setae
of hindcoxal pit not forming channel-like
structure. Male: maximum length of fla-
gellomere X 0.5-0.8 its width; maximum
length of flagellomere XI 2.2-3.6 times its
basal diameter; gonoforceps not expanded
exteroventrally (Fig. 2E, F) 2

2. Mesothorax without anteroventral process1;
mesopleural setae around scrobe in most
specimens longer than midocellar diameter
(shorter in some individuals). Female: clyp-
eal free margin sinuate between lobe and
orbit (Fig. 2A), hindcoxal pit oblong (Fig.

1 In occasional males the anteroventral mesothoracic process
is absent and the mesopleural setae are shorter than the mid-
ocellar diameter. Such specimens can be either figulus with
unusually short mesopleural setae, or minus without meso-
thoracic process. I cannot find characters for distinguishing
them.

2B-D) or (some specimens) circular, eva-
nescent in Japanese specimens

figulus (Linnaeus)

- Mesothorax with anteroventral process (Fig.
5B, C) in more than 95% of specimens;
mesopleural setae around scrobe shorter
than midocellar diameter. Female: clypeal
free margin almost straight between lobe
and orbit (Fig. 5A), hindcoxal pit circular
(Fig. 5D-F) minus (de Beaumont)

Trypoxylon figulus (Linnaeus)

Sphex figulus LINNAEUS, 1758:570. ! Holotype: 9, Sweden,
Uppsala (Linnean Society, London). — DAY, 1979:62. — In
Trypoxylon: LATREILLE, 1802:79; TSUNEKI, 1981:15 (rede-
scription, geographic variation). — In Apius: JURINE, 1807:
142.

Sphex fuliginosus SCOPOLI, 1763:292 (as fuliginosa, incorrect
original spelling). Holotype or syntypes: Carniolia (formerly
in Austria, since 1919 part of Italy and Yugoslavia), lost,
see ROGENHOFER UNO DALLA TORRE, 1882:599. ! Neotype:
2, Austria: "Carinthia, Ostkarawanken, Ebriach, 580-750
m, 21-29. VII. 1964, G. van Rossem," present designation
(CAS). — As probable synonym of figulus: VANDER LINDEN,
1829:42. — As synonym of figulus: subsequent authors.

Trypoxylon figulus var. majus KOHL, 1 883:657, 9, $ (as major,
incorrect original spelling). Holotype or syntypes: 9, Swit-
zerland: no specific locality (originally NHMV, Vienna, now
lost). ! Neotype: 2, Switzerland, "P. 3 VIII 84" and "Cn.
Tournier" (=Peney near Geneva, collection Tournier), pres-
ent designation (MHNG). New synonym. — DE BEAUMONT,
1945:477 (var. major); BLUTHGEN, 1951:234 (var. major);
DE BEAUMONT, 1958:206 (forma major), 1959:30 (same);
WOLF, 1959:15, 1 6 (figulus major); VALKEILA, 1961:244 (var.
major); DE BEAUMONT, 1964a:290, 1 964b:84 (forma major),
1965:56 (same), 1967:338 (same); BOHART AND MENKE, 1976:
346 (ssp. major); LOMHOLDT, 1976:267 (figulus major);
RICHARDS, 1980:45 (var. major).

Trypoxylon apicale W. Fox, 1891:1 42, 2 (as apicalis, incorrect

PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS

127

original spelling). Lectotype: 9, Canada (ANSP, Philadel-
phia), designated by CRESSON, 1928:52. — SANDHOUSE, 1940:
156 (apicale). Synonymized by PATE, 1943:16.

Trypoxylon figulus barbarum DE BEAUMONT, 1957: 9, $. Ho-
lotype: 3, Morocco: Marrakech (Mus. Zool. Lausanne). New
synonym. — BOHART AND MENKE, 1976:346.

Trypoxylon figulus yezo TSUNEKI, 1956:29, 9, $. Holotype: 2,
Japan: Hokkaido [=Yezo]: Jozankei (K. Tsuneki collection,
Mishima). New synonym. — BOHART AND MENKE, 1976:
346; TSUNEKJ, 1981:21 (summary of faunistic data).

Trypoxylon fieuzeti GINER MARI, 1959:389, 6. ! Holotype: 3,
Morocco: Fez (IEE, Madrid). Synonymized with figulus bar-
barum by SUAREZ, in GINER MARI', 1959:400.

COMMENTS ON NEW SYNONYMS. — T. figulus
barbarum was based mainly on the elongate male
flagellomere XI. Because flagellomere XI varies
in length (see Geographic Variation below) this
subspecies is not recognized here. I also feel that
a formal name for the Japanese populations (fi-
gulus yezo) is unwarranted on morphological or
other grounds.

COMMENTS ON NEOTYPES.— The identity of
Sphex fuliginosus has never been satisfactorily
established, because the original description is
inadequate and the original material is lost (Ro-
genhofer und Dalla Torre, 1882). Consequently,
the name can only be denned by designation of
a neotype. In selecting a specimen of Trypoxylon
figulus as a neotype of Sphex fuliginosus I have
followed the traditional interpretation of the last
name.

The original material of Trypoxylon majus
cannot be found in the Vienna Museum (Dr. M.
Fischer's letter of 21 October 1982) and must be
lost. However, this name indicates a large body
size, and figulus averages larger than either me-
dium or minus. A neotype of majus has also been
designated.

DIAGNOSIS. — Most specimens of figulus differ
from medium and minus in having the meso-
pleural setae around the scrobe slightly longer
than the midocellar diameter. However, the setae
length is slightly less than this diameter in some
specimens from southern France and the Iberian
Peninsula (as they are in the other two species).
The anteroventral mesothoracic carina is sin-
uate, curved posterad mesally, but unlike most
minus it has no process. The free margin of the
female clypeus (Fig. 2A) is usually sinuate be-
tween orbit and the median projection (free mar-
gin concave in medium, almost straight in mi-
nus). However, the free margin is almost evenly
concave in certain specimens from Spain (almost
like medium, which is unknown from Spain), in

a specimen from Zirbelwald, Austria, and one
from Balderschwang, Federal Germany. The fe-
male hindcoxal pit is mostly oblong (Fig. 2B-D)
in western palearctic specimens, but occasionally
it is nearly circular, as in medium and minus; it
is evanescent in Japanese females. In the male,
the maximum length of flagellomere X equals
0.65-0.8 of its width (the lowest ratios are ob-
served is specimens in which flagellomere XI is
short, and vice versa); the maximum length of
flagellomere XI usually is 2.4-3.6 times the basal
diameter instead of 2.0-2.2 in most medium, but
only 2.2 times in occasional specimens (which
differ from medium in having a longer meso-
pleural vestiture and a shorter flagellomere X).
Body length 9-12 mm in female, 7.5-10 mm in
male.

GEOGRAPHIC VARIATION. — In most males (in-
cluding the two males seen from Portugal), the
maximum length of flagellomere XI equals 2.4-
2.7 times its basal diameter, but in occasional
specimens it is only 2.2 times (e.g., in a male
from Wachseldornmoos, Switzerland); it is 2.7-
3.2 times its basal diameter in Spanish individ-
uals, and 3.3-3.6 times in Moroccan individuals.

LIFE HISTORY.— Many specimens of figulus
(voucher specimens examined by me) were reared
from nests established in wood (Wolf, 1959). Six
females and 1 7 males examined were reared by
O. Lomholdt from nests in reed stems which had
been used for thatching roofs at Tisvilde Hegn,
Denmark.

GEOGRAPHIC DISTRIBUTION (Figs. 3, 4).— Most
of the Palearctic Region between Great Britain
and Japan, and also eastern North America (east-
ern Canada and northeastern USA).

RECORDS (Old World). —Algeria ( 1 <3): El Harrach (as Maison
Carree, apical flagellomeres missing, BMNH).

Austria: (102 9, 50 6, NHMV if not indicated otherwise):
Karnten: Afritzer See (WJP), Ebene Reichenau (RMNH),
Ebriach in Ostkarawanken (WJP), Eisenkappel, Mallnitz (ZMB),
Mauthen (ZMB), NOtsch, Waidisch bei Ferlach (FSAG, JG).
Niederosterreich: Bisamberg near Vienna (NHMV, CU), Buck-
lige Welt S Vienna, Briihl, Dornbach (CAS, NHMV), Eichkogel
near Vienna (RMNH), Guntramsdorf (DEI), Hainbach (FSAG),
Hamburg an der Donau (ZMB), Herzogenburg, Herzograd (JG),
Kalksburg near Vienna, Krumbach, Lobau near Vienna
(NHMV, ZMH), Marchfeld (ZMB), Mistelbach (ZMH), Mo-
dling (ZMH), Oberweiden (DBB), Piesting, Purk (W Krems),
Rappendorf bei Molk (AWE), Roggendorf bei Melk (JG), Rohr
im Gebirge, Schneeberg, Stillfried (ZMH), Traismauer, Weid-
lingsbach (ZMH), Wien (NHMV, FSAG, ZMH) including
Donauauen, Kahlenberg and Turkenschanze. Oberosterreich:
Frauenstein (JG), Gemeinde Reichenthal (AWE), Gutau (FSAG,
JG), Hofkirchen (FSAG), Innerbreitenau (FSAG, JG), Kalten-

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FIGURE 2. Trypoxylon figulus: A— female clypeus, B— female hindcoxa ventrally, C— female hindcoxal pit, vertical view,
D— same, oblique view, E— male genitalia (arrow: area shown in F), F— same, portion of gonoforceps.

berg (AWE), Linz (HY, NHMV), Molln, MUhlviertel (JG),
Neumarkt (JG), Oberwallsee bei Miillachen (JG), Riedegg bei
Gallneukirchen (JG), Sankt Willibald (AWE), Ternberg (JG),
Welserheide, Zeissberg bei Freistadt (FSAG, JG). Salzburg:
Koppl bei Aschach (JG), Salzburg (BMNH). Steiermark: Leut-
schach (JG), Sankt Ulrich (JG), Tragoss-Oberort (DBB), Wein-
burg (FSAG). Tirol: Huben in eastern Tirol (ZMB), Innsbruck
(NHMV, ZMB), Iselsberg (DBB), Lienz (RMNH), Obladis,
Hopfgarten, Salvenberg (CAS), Zirbelwald near Obergurgl, 1
km SW Zwiselstein in Otztal. Voralberg: Ittensberg.

Belgium (95 9, 63 $, FSAG if not indicated otherwise): Bra-
bant: Evere, Genval, Gistoux, Grez-Doiceau, Mont-Saint-Gui-
bert, Nethen, Rhode-Sainte Agathe, Thorembais-Saint-Trond,
Uccle, Waterloo. Hainaut: Aiseau, Athis, Barry, Binche, Bous-
su, Bouvignies, Fleurus, Orcq, Seneffe, Taintignies, Velaines,

Wanfercee. Liege: Acosse, Aubel, Barchon, Ben-Ahin, Beyne:
ca 15 km SE Liege (BMNH), Beyne-Heusay, Cerexhe, Char-
neux, Chevron, Clermont-sur-Berwinne, Fleron, Foret de
Grunhault, Francorchamps, Henri-Chapelle, Hombourg, Ju-
pille, La Calamine, La Reid, Lontzen, Montzen, Pepinster,
Queue-du-Bois, Romsee, Spa, Xhendelesse, Welkenraedt.
Limburg: Berg pres de Tongres, Bocholt, Godsheide, Tongres.
Luxembourg: Amonines, Hotton, Les Epioux, Lomprez, Ozo,
Saint-Medard, Sampont, Smuid, Waharday, Wibrin. Namur:
Aische, Alle (RMNH), Andenne, Baillonville, Belgrade, Bievre,
Branchon, Champion, Eghezee, Ernage, Feschaux, Gembloux,
Gesves, Grand Leez, Ham-sur-Sambre, Lonzee, Mount-Gau-
thier, Saint-Aubin, Saint-Gerard, Saint-Marc, Sorinnes, Sau-
veniere, SombrefTe, Winenne.
Bulgaria (1 $): Rila Mts. (DEI).

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129

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 10

FIGURE 4. Trypoxylon figulus: geographic distribution in the New World.

Czechoslovakia (1 2, 3 3): Jihomoravsky Kraj: Vranov W
Znojmo (as Frain, NHMV). Vapadoslovensky Kraj: Sturovo
(SMT).

Denmark (41 9, 33 6, ZMK if not indicated otherwise): Al-
borg: Vegger. Bornholm: Bastemose. Frederiksborg: Asserbo,
Hillerod, Hundested, Jaegerspris Nordskov (tip of Horn-
sherred peninsula), Sorte Mose near Farum, Store Karlsminde
near Lynaes, Tisvilde Hegn (FSAG, ZMK). Holbsek: Kongsore.
Maribo: Maribo. Kgbenhavn: Holte. Odense: vCbelo. Randers:
Glatved Strand on Djursland Peninsula, Mols Bjerge. Ring-
kgbing: Gindeskov, Kjelstrup (E Skjern). Sdnderborg: S0nder-
borg. Sor«: AgersO. Svenborg: Langeland Island: Hellenor.
lander: Rome (VH), Stensbaek Plantage. Vejle: Klattrup.

Finland: (47 9, 48 <3, HY unless indicated otherwise): Abo:
Lohja, PerniO (AKM), Ryma'ttyla (AKM), Turku (AKM).
Alandia: Ecker6(AKM, HY), Finstrom (AKM), Hammarland,
Jomala (AKM), Saltvik (AKM). Karelia Borealis: PyhSselka
(Hammaslahti). Nylandia: Helsinki (CU, HY), Parvoo = Bor-
ga, Pernaja. Ostrobotnia Australis: Koivulahti. Satakunta:
Loimaa (AKM), Yla'ne (AKM). Tavastia Australis: Hattula,
Hameenlinna, Janakkala, Nastola (AKM), PalkSne (AKM, HY),
Somero (AKM), Urjala, Vanaja, Yloja'rvi.

France (35 9, 16 <3): Alpes-Maritimes: Guillaumes (FSAG).
Ariege: Ax-les-Thermes (ZMB). Basses-Alpes: Allos (FSAG),
Annot (FSAG), Fugeret (FSAG), Les Dourbes (KMG), Mon-

tagane de Lure (ZMK), Peyresq (FSAG), Saint-Andre-les-Alpes
(FSAG). Bouche-du-Rhone: Marseille (FIS). Calvados: Lisieux
(FSAG). Corse: Corte (KMG). C6tes-du-Nord: Saint-Rieul.
Haute-Savoie: Dent d'Oche (MHNG), Mont Jorat (RMNH),
Val de Charmy (RMNH). Haute-Vienne: Rochechouart (FSAG).
Jura: Arbois. Loire-et-Cher: Blois (FSAG). Loire-Atlantique:
Foret de la Roche Bernard (RMNH), Herbignac (RMNH).
Saone-et-Loire: Uchizy (FSAG). Seine-et-Oise: Poissy (IEE).
Van Frejus (KMG), Gonfaron (FSAG), Montouroux (RMNH).
Vaucluse: Carpentras (RMNH). Yonne: Foissy-sur-Vanne
(FSAG).

Germany, Democratic (81 2, 29 $, DEI if not indicated oth-
erwise): Berlin: Berlin (DEI, HD, ZMB, ZSM). Cottbus: Alt
Dobern (ZMB), Muskau, Neu Zauche (ZMB), Schlieben (ZMB).
Dresden: Daubitz (SMT), Gersdorf near Kamenz (SMT). Er-
furt: Erfurt (CU), Gotha (ZSM). Frankfurt: Biesental, Ebers-
walde area. Gera: Blankenburg (ZMB), Jena (NHMV, ZMB).
Halle: Gernrode (ZMH), Halle (DEI, ZMB), KyftTiauser, See-
burg, Naumburg (TMB). Leipzig: Winkelmiihle. Magdeburg:
Arendsee (SMT). Neubrandenburg: Faule Ort, Naturschutz-
gebiet Muritzhof. Potsdam: Furstenberg (TMB), Zechlin (ZMB),
Zootzen. Rostock: Prerow, Rostock, Stralsund (DEI, ZMB).
Riigen: Hiddensee Island (DEI, SMT), Riigen Island: Monch-
gut (SMT) and Ummanz. Schwerin: Campow (ZMB), Schwerin
(ZMB), Wendeltorf near Schwerin.

PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM. AND MINUS

131

Germany, Federal (93 2, 51 3): Baden-Wiirtemberg: Enz-
klosterle (KS), Heidelberg (ZSM), Hochwacht (HW), Isny
(NHMB), Kaiserstuhl (ZMB), Karlsruhe (KS, ZMH), Kiissa-
berg (KS), Radolfzell (ZMH), Schwarzwald (SMT), Tiengen in
Wutach Valley (KS). Bayern: Abensberg (ZSM), Allach (ZSM),
Aschaffenburg (FIS), Balderschwang (KS), Bamberg (ZSM),
Ebenhausen (ZSM), Erdweg (ZSM), Horgersthausen near
Moosburg (ZSM), Ingolstadt (ZSM), Kahl (FIS), Miinchen
(FSAG, ZSM), Nurnberg(ZSM), Rotwand area (ZSM), Schlier-
see (ZSM), Tegernsee (ZSM). Hamburg (ZMH): Ochsenwarder,
Warwisch. Hessen: Battenfeld near Biederkopf (ZMH), Griln-
dau E Frankfurt (HW, HY), Marburg (HW, WJP). Nieder-
sachsen: 2 km NW Dotlingen (VH), Dorpen: 14 km SW Pa-
penburg (VH), Elbe Islands (VH), 5 km S Oldenburg (VH),
Pevestorf: 72 km SE Lauenburg (VH), Wobeck (ZMH). Nord-
rhein-Westfalen: Ahaus (ZSM), Neheim (FSAG), Leverkusen
(ZMH), Plettenberg(HW), Siegen (HW). Rheinland-Pfalz: low-
er Ahr valley (FIS), Mainz (KS), Nattenheim (FSAG), Worms
(FIS). Schleswig-Holstein: Amrum Island (VH), Eutin (KS),
Ihlkathe 2 km SE Kiel (VH), Lutjenburg (KS), Ratzeburg (ZMH),
Schierensee SW Kiel (VH), Schleswig (VH).

Great Britain (92 2, 106 3; BMNH unless stated otherwise):
Berkshire: Reading. Buckingham: Iver, Slough. Devon: Paign-
ton. Dorset: Wareham. Essex: Brentwood, Colchester, Epping
Forest. Gloucester: Chalford. Hampshire: Brockenhurst, Fleet,
New Forest, Wickham. Isle of Wight: Sandown, Shanklin. Kent:
Cobham, Darenth, Faversham (WJP), Goudhurst. London:
Hampstead, Mill Hill, Mitcham Common, Norwood, Putney.
Northampton: Ashton Wold (Oundle). Oxford: Goring, Ox-
ford, Tubney near Oxford. Somerset: Dunster. Suffolk: Arger
Fen, Bury St. Edmunds, Dunwich. Surrey: Byfleet, Esher, Hor-
sell, Weybridge. Sussex: Midhurst: Ambersham Common.

Greece (2 2, 4 3): Peloponnesus (de Beaumont, 1965): Mega
Spilaion, Pirgos, and Taygetus. Sterea Ellas: Karpenission
(KMG). Thessalia: Aspropotamos near Kalabaka (KMG).

Hungary (13 2, 3 <3): Bacs-Kiskun: Kalocsa (TMB), Tabdi
(TMB). Gyor-Sopron: Neusiedlersee (NHMV). Somogy: Bal-
atonszemes (TMB). Szolnok: Jaszbereny (TMB). Tolna: Si-
montornya (NHMV). Veszprem: Tihany Peninsula on Balaton
Lake (HD, TMB, WJP).

Italy (19 2, 8 3): Emilia- Romagna: Cattolica (RMNH). Lom-
bardia: Pavia: Cignolo Po (MSNM), Sondrio: Valtellina (KS).
Piemonte: Alpignano (GP), Colle di Sestriere in Alpi Cozie
(GP), Murazzano (GP), San Benedetto Belbo 20 km S Alba
(GP), Val d'Angrogna in Alpi Cozie (WJP). Valle d'Aosta:
Bresson near St. Vincent (GP). Venezia Giulia: Trieste (CU,
NHMV). Trentino-Alto Adige: Bolzano (as Bozen, NHMV),
Cavalese (BB), Collalbo (de Beaumont, 1959), Ortisei (NRS),
Trafoi (NHMV).

Japan (Tsuneki 1981): western Hokkaido (Esashi, Hakodate,
Jozankei, Kamikawa) and central Hondo (Prefectures: Fukui,
Ishikawa, Kyoto, Nagono, Niigata, Saitama, and Yamanashi).
Specimens studied: 4 2, 4 <?; KT, TN.

Morocco (1 2, 2 3): Asni, 1250 m alt. (paratype 2 offigulus
barbarum, RMNH), Fez (holotype offieuzeti, IEE), Marrakech
(BMNH).

Netherlands (108 2, 73 3, RMNH if not indicated otherwise):
Drenthe: Emmen, Erm, Havelte, Sellingen, Wijster, Zuidlaren.
Gelderland: Apeldoorn (FSAG), Barneveld, Hulshorst, Nij-
kerk, Putten, Renkum, Vierhouten, Wageningen (GVR,
RMNH). Groningen: Onnen. Limburg: Epen (GVR), Haelen,
Heerlen, Helden, Posterholt. Noord Brabant: Empel, Helen-
aveen, Neerijnen, Udenhout, Ulvenhout. Noord Holland: Aer-
denhout, Hilversum, Laren. Overijssel: Heino, Wezepe. Utrecht:

Baarn, Bilthoven, de Bilt, den Dolder. Zuid Holland: Delft
(GVR), Hillegersberg, Leiden, Leiderdorp, Loosduinen, Oeg-
stgeest, Rijnsburg, Rotterdam, 's Gravenhage, Warmond.

Norway ( 1 2, 1 3): Buskerud: Al (as Aal, DEI). Oppland: Lom
(RMNH).

Poland (35 2, 24 3): Biarystok: Biafowieza (as Bialowies, FIS,
ZMB), Lenkowo near Grajewo (ZMB). Bydgoszcz: Bydgoszcz
(as Bromberg, SMT). Kielce: Gory Pieprzowe near Sandomierz
(CAS), Kielce (TMB), Sandomierz (CAS). Lublin: Kazimierz
on Wisla River (WJP). Warszawa: Podkowa Lesna (CAS).
Wroclaw: Muszkowice near Henrykow (WJP), Osola: 25 km
NW Wroclaw (WJP), Wroclaw (CAS, WJP). Szczecin: Bielinek
(as Bellinchen, ZMB), Miedzyzdroje (as Misdroy, ZMB). Zie-
lona Gora: Kostrzyfi (as Custrin, ZMB).

Portugal (2 2, 2 3): Douro: Resende (RMNH). Estramadura:
Lisboa (RMNH).

Romania (8 2, 9 3, MGA if not indicated otherwise): Bacau:
Lacul Bicaz 20 km W Piatra Neam{ (HD). Brasov: Sighisoara,
Turnisor near Sibiu. Bucuresti: Branesti (WJP), Budesti. Con-
stanta: Crisan, Hagieni. Crisana: Ineu (as BorosjenO, TMB).
Orasul Bucuresti: Mogosoaia, Pantelimon. Timisoara: Esel-
nija, Jupalnic near Mehedintj, Mehadia (NHMV), Orsova
(RMNH).

Spain (53 2, 5 1 3): Alicante: Alicante (KMG), Orihuela (IEE).
Avila (SFG): Becedas, Gilbuena, Puerto Castilla, Solana de
Bejar, Tremedal. Barcelona: Canet de Mar (FSAG). Caceres
(SFG): Baftos de Montemayor, Cabezuela del Valle, Gargan-
tilla, La Garganta. Cadiz: Algeciras (NHMV). Ciudad Real:
Ruidera (FJS). Granada: Salobrefla (VH). Huesca: Arguis
(RMNH), Torla (FSAG), Valle de Ordesa. Lerida: Bohi, Lago
di San Mauricio (FSAG). Logrono: San Roman de Cameros
(IEE). Madrid: El Escorial (IEE), Sierra de Guadarrama (IEE).
Salamanca (SFG): Bejar, Cantagallo, Colmenar de Monte-
mayor, El Cerro, La Cabeza de Bejar, Lagunilla, Montemayor
del Rio, Navacarros, Palomaresde Bejar, Peflacaballera, Pueb-
la de San Medel, San Medel, Sorihuela, Valdehijaderos, Val-
lejera de Rio Frio. Teruel: Albarracin (KMG, ZSM), Libros
(FSAG). Toledo: Toledo (RMNH, WJP). Valencia: La Eliana
(FSAG). Valladolid: Olmedo (FJS, WJP), Simancas (RMNH,
WJP), Valladolid (FJS, WJP), Villa Baflez (FJS), Villalba de
los Alcores (FJS). Zamora: Montamarta (FSAG).

Sweden (43 2, 403, NRS if not indicated otherwise): Dalarna:
Ta'ktbo. Oland: Arontorp, Glomminge, Hogsrum, Halltorps
hage, Morbylanga, Repplinge, Vickleby. Ostergotland: St. Anna
Korsna's, Simonstorp. Skane: Ahus, Halsingborg, HavSng, H68r
distr. (BMNH), Torekov (ZMK), Trolleholm, Vitemolla. Sma-
land: Hagby, Jonkoping, Kalmar, Kalmar Sund (Bla Jungfrun),
Soderakra. Sodermanland: Haggenas, Ma'larhojden, Morto,
Tullgarn, TyresS, Tyreso - Brevik. Uppland: BjorkO, Habo -
Tibbie, Osteraker, Radmanso, Svartsjo, Varmdo, Vassunda.

Switzerland (78 2, 41 3, NHMB if not indicated otherwise):
Bern: Adelboden, Bantiger (7 km NE Bern), Batterkinden, Bern,
Biischiwald forest (5 km SSW Bern), Gadmen, Miirren, Signau,
Spiez, Wachseldommoos bog (27 km SE Bern), Zweisimmen,
Zwischenfluh (16 km SSW Thun). Geneve: Geneve (MHNG,
NHMV), Genthod (NHMV), Peney (MHNG, NHMV). Grau-
biinden: Chur (ZSM), Pare National Suisse (MHNG), Somvix,
Versam. Luzern: Entlebuch Graben (CAS). Sankt Gallen: Vat-
tis. Valais: Ausserberg ( 1 1 km W Brig), Ayel - Zinal (RMNH),
Berisal (BMNH, NHMB, NHMV), Chalet a Gobet (BMNH),
Champery, Eusegne (as Usegne), Evolene, Hauderes, Inden,
Lotschental, Martigny, Riederalp (8 km NNE Brig), Saas, Sankt
Niklaus, Sierre (NHMV), Simplon (BMNH), Stalden, Verbier
(BMNH), Vissoye, Zermatt. Vaud: Nyon (MHNG).

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Turkey (6 9, 3 3): Afyon: Cay (FSAG). Ankara: Ankara (as
Angora, TMB), Karagol (BMNH). Denizli: SaraykOy (as Seraj-
Koj, TMB). Istanbul: Sile area (BMNH). Kenya: Konya (JG).
Kiitahya: Karat Dagi (BMNH). Samsun: Koprubasi (de Beau-
mont, 1967). Tokat: Arguslu above Niksar(BMNH). Trabzun:
Zigana Dagi (de Beaumont, 1967).

USSR: Armenian SSR: Yerevan area (8 S, ZMMU). Azer-
baydzhan SSR (1 2): Khanlar (as Helenendorf, NHMV). Ka-
zakh SSR: Vostochnokazakhstanskaya obi.: foothills of Azu-
tau Khrebet 18 km N Alekseyevka (1 3, VLK); Kalbinskiy
Khrebet, 20 km SE Leninka (1 9, VLK); 15 km NW Ust'-
Kamenogorsk (1 3, VLK). Latvian SSR (2 9, 8 S): Wezkukkul,
20 km S Jaunjelgava (ZMB). Lithuanian SSR (19,2 S): Ignalina
(DEI). Russian SSR: Belgorodskaya oblast: Valuyki (2 2,
NHMV). Chelabinskaya oblast: Ilmenskiy Zapovednik (1 9,
ZMMU). Gorno-Altayskaya Avtonomnaya oblast: Uymen
River (2 S, ZMMU). Ivanovskaya oblast: Kineshma (1 2,
ZMMU). Karelian ASSR (2 2, 1 S, HY): Salmi, Sortavala.
Kuybyshevskaya oblast: Mirnyi (2 2, VLK). Leningradskaya
oblast: Solnechnoye (as Ollila, 1 6, HY), Streltsovo (as Muola,
1 2, HY), Suursari Island (as Hogland, 1 3, HY). Karachayevo-
Cherkesskaya oblast: Teberda-Dzhamagat (2 9, HD). Moskov-
skaya oblast: Krylatskoe near Moskva (4 2, 1 3, ZMMU), Mosk-
va (1 2, ZMMU), Mytishchi (1 3, ZMMU), Zavety Il'icha
(1 3, ZMMU). Orenburgskaya oblast: Kargala near Orenburg
(1 2, ZMB). Ukrainian SSR: Otuzy Valley in Crimea (2 9, 2 3,
ZMMU), Podgortse E Lvov (as Podhorce, 1 2, NHMV), Se-
bastopol area (1 3, ZMMU).

Yugoslavia ( 1 1 2, 6 3): Croatia: Plitvice (RMNH), Senj (VH).
Kosovo: Pet (TMB). Macedonia: Mavrovska Valley (RMNH),
Ohrid - Resen (RMNH). Slovenia: Begunje near Postojna (KS),
Kranjska Gora (as Weissenfels, NHMV), Portoroz (FIS), Rad-
enci in Mura Valley (near Austrian border, as Radein, NHMV),
Vipava (as Wippach, NHMV).

RECORDS (New World) (Krombein 1951:955, 1979:1643;
Finnamore 1982:1 15). -Quebec: Duchesnay (1 2, USNM), He
Perrot (2 2, LEM), Lakeside (1 2, LEM), Levis (3 2, LEM),
Montreal (1 2, CU; 2 3, LEM), Ste. Anne de Bellevue (3 2, 6
3, LEM), Ste. Annes (1 2, 3 3, LEM), St. Hilaire (5 2, LEM),
St. John's Co. (1 2, LEM).

Ontario: Ottawa (2 9, 2 3, CNC), St. Anne's (4 2, CNC).

Maine: Bangor (1 2, MCZ).

New Hampshire: Glen House (1 3, USNM).

Massachusetts: Cambridge (1 2, USNM), Dorchester (1 2,
MCZ), Fall River (1 2, USNM), no specific locality (1 9, WSU).

Trypoxylon minus de Beaumont

Trypoxylon figulus var. minus DE BEAUMONT, 1945:478, 2, 3
(as minor, incorrect original spelling). Holotype: 2, Switzer-
land: Cologny near Geneva (Mus. Zool. Lausanne). —
BLUTHGEN, 195 1:234 (var. minor); DE BEAUMONT, 1958:206
(forma minor), 1959:30 (same); Wolf, 1959:15, 16 (figulus
minus); VALKEILA, 1961:144 (var. minor); DE BEAUMONT,
1964a:290, 1964b:84 (forma minor), 1965:56 (same), 1967:
338 (same); BOHART AND MENKE, 1976:346 (ssp. minor);
RICHARDS, 1980:45 (var. minor).

Trypoxylon figulus koma TSUNEKI, 1956:28, 9, 3. Holotype: 9,
Korea: Mt. Kodai (K. Tsuneki collection, Mishima). New
synonym.— BOHART AND MENKE, 1976:346; TSUNEKI, 1981:
20.

Trypoxylon figulus medium: WOLF, 1959:fig. b.

DIAGNOSIS. — Most minus can be recognized
by the presence of a flat, median process emerg-

ing from the anteroventral mesothoracic carina
(Fig. 5B, C). However, the process is absent in a
female and two males from Linz, Austria, and
also one of the females from Gonfarons, France,
which otherwise do not differ from the remaining
minus examined. Unlike most figulus, the me-
sopleural setae around scrobe are shorter than
the midocellar diameter. In the female, the me-
dian clypeal projection is shorter than in medi-
um; unlike figulus and medium, the clypeal free
margin is scarcely sinuate or evenly arcuate be-
tween the orbit and the projection (Fig. 5A). The
female hindcoxal pit is circular or nearly so (Fig.
5D-F), unlike most figulus. The maximum length
of male flagellomere X is 0.5-0.7 times its width;
flagellomere XI is longer than in most medium:
its maximum length equals 2.2-3. 1 times its bas-
al diameter. Body length 6-9 mm in female,
5-7.5 mm in male.

VARIATION. — In most females, the free margin
of the clypeal projection is emarginate, but it is
entire in a female from Finland, one from France
and one from Sweden.

In most males, the maximum length of fla-
gellomere X is 0.6-0.7 times its width, and the
maximum length of flagellomere XI is 2.2-2.4
times the basal diameter. These ratios are 0.8
and 2.6, respectively, in some males from Bel-
gium, 0.6 and 2.6 in the single male examined
from Sardinia, 0.8 and 3.0 in the single male
from Corsica, and 0.5 and 3.1 in the single male
from Gerona, Spain.

LIFE HISTORY.— A female from Elender Wald,
Austria (NHMV), was reared from a gall of An-
dricus kollari (Hartig). I was unable to find this
locality.

GEOGRAPHIC DISTRIBUTION (Fig. 5). — Palearc-
tic Region between western Europe and Korea,
north to about 64°N in Norway and Sweden and
about 67°N in Finland, south to northern Med-
iterranean countries including Sardinia and Sic-
ily, northern Turkey and southern Kazakh SSR.
Unknown from Great Britain and North Africa.
The absence of this species in Great Britain sug-
gests that it reached northwestern Europe only
recently, after separation of the British Isles from
the continent.

RECORDS.— Austria (41 2, 52 3, NHMV if not indicated oth-
erwise): Burgenland: Breitenbrunn (JG), Donnerskirchen on
Neusiedlersee (KS), Mogersdorf (JG), Morbisch: 5 km S Rust
(RMNH). Karnten: Bleiberg (RMNH), Mallnitz (ZMB), Wai-
disch bei Ferlach (FSAG). Niederosterreich: Anninger, Bis-
amberg near Vienna, Briihl, Dornbach, Hainbach, Hainsburg

PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS

133

FIGURE 5. Trypoxon minus: A— female clypeus, B— mesothoracic venter, C—antero ventral mesothoracic process, D—
female hindcoxa ventrally. D— female hindcoxal pit, vertical view, E— same, oblique view.

(ZMB), Kahlenberg near Vienna, Neunkirchen (FSAG, JG),
Piesting, Purgstall (JG), Schneeberg, Traismauer, Wien (Pra-
ter). Oberosterreich: Almsee (NHMV), Gutau (JG), Inner-
breitenau (JG), Kremsmiinster (FSAG), Linz (HY, JG, NHMV),
Rutzing bei Horsching (JG), Sarleinsbach, Zeissberg bei Frei-
stadt (FSAG, JG), Zellhof bei Bad Zell (JG). Salzburg: Salzburg
(BMNH, FSAG). Steiermark: Admont (FSAG), Gleichenberg
(JG), Riegersburg (JG), TragOss-Oberort (DBB), Weinburg (JG).
Tirol: Huben (ZMB), Innsbruck, Kals (RMNH), Stubai (WJP),
Wenns (BMNH), Zwieselstein in Otztal.

Belgium (348 2, 379 <5, FSAG if not indicated otherwise):
Brabant: Berchem-Sainte-Agathe, Groenendael, Monstreux,
Rhode-Saint-Genese, Rixensart, Tilly, 't Roth (RMNH), Wo-
luwe-Saint-Pierre. Hainaut: Gilly, Gosselies, Lobbes-Calvaire,
Quaregnon. Liege: Bleyberg, Chaudfontaine, Clermont-sur-
Berwinne, Dalhem, Embourg, Esneux, Foret de Grunhault,
Fouron-Saint-Pierre, Hergenrath, Herstal, Jupille, Liege, Queue-
du-Bois, Remersdael, Spa, Xhendelesse, Wandre. Limburg:
Bassenge, Eben, Kanne, Sint Pietersberg (RMNH), Tongres,
Wonck. Luxembourg: Bodange, Chatillon, Les Epioux. Namur:

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135

Beauraing, Couvin, Felenne, Gembloux, Gesves, Jambes, La
Plante, Malonne, Mont-Gauthier, Namur, Saint Marc, Wil-
lerzie, Yvoir. West-Vlaanderen: Sint Pieter.

Bulgaria (1 3): Chepalare Pass in Rhodope Mts. (ZMT).

Czechoslovakia (4 9, 4 3): Jihomoravsky Kraj: Vranov W
Znojmo (as Frain, NHMV). Vapadoslovensky Kraj: Sturovo
(HD, SMT).

Denmark (2 2, ZMK): Maribo: Lolland Island: Roden Skov.
Vejle: Klattrup.

Finland (105 9, 124 3, HY if not indicated otherwise): Abo:
PerniO (AKM), Rymattyla (AKM), Sarkisalo (AKM), Turku
(AKM). Alandia: EckerO (AKM, HY, ZMK), Finstrom (AKM),
Geta (ZMK), Hammarland, Lemland. Karelia Australis: Vi-
rolahti. Karelia Borealis: Herajoki on Pielinen Lake (ZMK),
Kesalahti, Kitee (AKM, HY), Polvijarvi (AKM), Tohmajarvi.
Karelia Ladogensis: Parikkala, Simpele. Kuusamo: Pera Posio.
Lapponia Kemensis: Sodankyla. Nylandia: Helsinki, Pernaja
(as Parna). Ostrobottnia Australis: Koivulahti. Ostrobottnia
Borealis: Ra'vasjarvi, Rovaniemi. Ostrobottnia Media: Siika-
joki. Satakunta: Loimaa (AKM), Suoniemi. Savonia Australis:
Lappeenranta, Rautjarvi. Tavastia Australis: Aitolahti, H3-
meenlinna, Hattula, Janakkala, Kalvola, Kangasala, Lahti
(AKM), Lammi, Luopioinen (AKM), PaMne, Pirkkala (as
Birkkala), Somero (AKM), Tyrvanto, Vanaja, Ylojarvi.

France (24 9, 17 3): Basses-Alpes: Allos (FSAG), Annot
(FSAG), Colmars (FSAG), La Javie (FSAG), Peyresq (FSAG),
Saint-Andre-les-Alpes (FSAG). Corse: Bonifacio (KMG), Col
de Sorba (KMG). Drome: Dieulefit (RMNH). Haute Savoie:
Chamonix (RMNH). Haut-Rhin: Rouffach (FSAG). Loiret:
Lorris (RMNH). Lozere: Sainte Enimie (BMNH). Nievre:
Nevers (RMNH). Puy-de-D6me: Beaune (FSAG, ZMK), Besse
(BMNH). Pyrenees-Orientales: Vinca (VH). Van Gonfarons
(FSAG).

Germany, Democratic (33 2, 20 3, DEI if not indicated oth-
erwise): Berlin: Berlin-Borgsdorf (HD). Dresden: Freital (SMT),
Gersdorf near Kamenz (SMT), Moritzburg (SMT). Erfurt:
Nordhausenarea(HD). Frankfurt: Eberswalde area (DEI, HD),
20 km NW EisenhUttenstadt (HD), Schonwalde near Berlin
(HD). Gera: Beutnitz bei Jena. Halle: Halle (DEI, ZMB),
Kyffhauser (DEI, ZMB). Karl-Marx-Stadt: Freiberg. Leipzig:
Dornreichenbach near Wurzen, Leipzig, Reglitz bei Schkeu-
ditz. Magdeburg: Haldensleben, Stapelburg (ZMH). Neubran-
denburg: Serrahn near Neustrelitz (HD), Teterower See (HD).
Rostock: Rostocker Heide. Schwerin: Perleberg, Schwann near
Gilstrow.

Germany, Federal (59 2, 42 3): Baden-Wiirttemberg: Enz-
klosterle (KS), GrOtzingen (KS), Heidelberg (BMNH), Kai-
serberg (ZMH), Kaiserstuhl (ZMB), Karlsruhe (KS, ZMH),
Kiissaberg (KS), Tubingen (KS), Windenreute near Emmen-
dingen (ZMB). Bayern: Aschaffenburg (VH), Balderschwang
(KS), Karlstadt am Main (KS), Lohr am Main (KS), Main-
franken (FIS), Miltenberg (FIS), Munchen (ZSM), Oberau in
foothill of Ammer Mts. (KMG), 32 km S Wurzburg (RMNH).
Hessen: Dillenburg (HW, WJP), Fulda (VH), Marburg (HW,
WJP), Taunus (KS). Rheinland-Pfalz: Bad MUnster am Stein
(KS), Burgen an der Mosel (TMB), Donnersberg, Mainz (KS).
Niedersachsen: Sage: 25 km S Oldenburg (VH). Schleswig Hoi-
stein: LUtjenburg (KS), Russee near Kiel (VH), Schierensee SW
Kiel (VH). Wesrfalen: Plettenberg (HW), Siegen (HW).

Greece (2 2, 2 3): Ionian Islands: Isle de Levkas (as Lefkas,
BMNH). Sterea Ellas: Timfristos (KMG).

Hungary (1 2, 1 <J): Balaton (HD).

Italy (23 2, 143): Abruzzi: Scanno (KMG). Emilia-Romagna:
Bologna: Pracchia (RMNH). Ronzano (NHMV); Modena:

Zocca: Montetortore (NHMV). Friuli-Venezia Giulia: Ge-
mona (JG), Prosecco (NHMV), Tarvisio (NHMV). Lombardia:
Varese: Mercallo (MSNM). Piemonte: Borgomale (GP), Cas-
telnuovo in Asti (GP), Chianale (GP), Condove (GP), San
Benedetto Belbo ca 20 km S Alba (GP), Val d'Angrogna in
Alpi Cozie (1 2, del. J. de Beaumont, WJP), Vinovo 10 km S
Torino (GP). Sardegna: Aritzo (BMNH), Desulo (BMNH).
Sicilia: Piano Battaglia (KMG). Trentino-Alto Adige: Colle
Isarco (MSNM), Merano (as Meran, FSAG). Valle d'Aosta:
Buthier (ZMB). Veneto: Padova (de Beaumont, 1959). Venezia
Giulia: Trieste (NHMV).

Korea: North Korea: Mt. Kodai near 38th parallel (Tsuneki,
1956, 1981), Mt. Sombo near 38th parallel (1 9, 1 3). South
Korea: Seoul (Tsuneki, 1956, 1981).

Netherlands (5 9, 9 3, RMNH unless indicated otherwise):
Limburg: Elkenrade, Epen, Maastricht, Slenaken, Vijlener Bosch
(GVR). Noord Brabant: Empel. Zuid Holland: Leiden.

Norway (1 1 2, 19 3): Akerhus: Oslo (as Kristiania, ZMUB).
Hordaland: Granvin (ZMUB). Nord Trandelag: Rora (FSAG).
Oppland: Lorn (RMNH). Sogn og Fjordane: Balestrand
(ZMBU). Vestagder: Kristiansand (ZMUB). Vestfold: Herstad
(ZMUB).

Poland (7 9, 1 3 3): Biatystok: Bialowieia (as Bialowies, ZMB).
Krakow: Ojc6w (SMT), Pieniny Mts. (WJP). Wroclaw: Kar-
konosze Mts. (as Riesengebirge, ZMB), Osola: 25 km NW
Wroclaw (WJP), Roscislawice (WJP), Sobotka: 35 km S
Wroclaw (WJP), Wroclaw (CAS), 10 km NE Wroclaw (WJP).
Rzeszow: Jaroslaw (CAS).

Portugal (1 9): Douro: Resende (RMNH).

Romania (6 9, 3 3, MGA if not indicated otherwise): Bu-
curesti: Ciolpani, Peris., Videle (WJP). Constanta: Babadag.
Ploesti: Valea Longa-Gorgota. Timisoara: Es.elni{a, Ogradina,
Or§ova.

Spain (2 2, 4 3): Gerona: Gerona (BMNH). Huesca: Sierra
de Oroel (FSAG), Valle de Ordesa (FIS). Salamanca: Horcajo
de Montemayor (SFG), Valdehijaderos (SFG).

Sweden (34 9, 44 3, NRS if not indicated otherwise): Bo-
huslan: Ljung. Dalarna: Ludvika, Nas, Stora Kopparberg,
Taktbo. Gastrikland: Sandviken. Gotland: Fardume Tr8sk E
Larbro (ZMK), Visby, St. KarlsO. Jamtland: Undersaker. Narke:
Orebro. Oland: Halltorps hage, HOgsrum, Rapplinge. Skane:
HOOr distr. (BMNH), Skaralid, northern Skane (BMNH). Os-
t ergot land: Kvarsebo, Norrkoping, Simonstorp. Smaland: Bar-
keryd, Horeda, jarsnas, Ljungarum. Sodermanland: Flisby,
Runmaro, Tullgarn, Tyreso - Svartocken. Uppland: Harparbol,
Radmanso, Stockholm, Uppsala (HY), Vassunda, Vira bruk.
Vasterbotten: Handene. Vastmanland: Sala.

Switzerland (21 2, 26 3, NHMB if not indicated otherwise):
Bern: Batterkinden, Bern, Burgdorf, Grauholz forest (up to 9
km NNE Bern), Wachseldornmoos bog (27 km SE Bern). En-
gadin: Zuoz (AWE). Geneve: Geneve Cologny (BMNH), Gen-
thod (NHMV), Peney (MHNG). Graubunden: Pare National
Suisse (de Beaumont, 1958), Somvix. Tessin: Lugano (FSAG).
Valais: Binntal (15 km NE Brig), Grimentz (FSAG, RMNH),
St. Luc (RMNH), Simplon (BMNH), Zermatt (RMNH). Vaud:
Nyon (MHNG). Zurich: Wadenswil.

Turkey (1 9, 2 3): Artvin: Artvin, Berta (BMNH). Istanbul:
Belgrat Orman (de Beaumont, 1967). Samsun: Bafra (BMNH).

USSR: Georgian SSR: Bakuriani (1 9, ZMMU). Kazakh SSR:
Chimkent, 2000 m ( 1 3, WJP); Razdolnyi area, 50°43'N, 8 1°06'E
( 1 3, VLK), Zapovednik Aksu-Dzhabagly ( 1 2, ZMMU). Kirgh-
iz SSR: Arkit in Chatkalskiy Khrebet (1 3, ZMMU). Russian
SSR: Karelian ASSR: ozero Tumas (as Kolatselka, 1 2, HY).
Kemerovskaya oblast: 60 km SE Novokuznetsk (1 3, VLK).

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Leningradskaya oblast: Kondratyevo W Vyborg (as Sa'kkijarvi,
1 2, HY). Moskovskaya oblast: Mytishchi (1 6, ZMMU). Perm-
skaya oblast: Nizhnyaya Kurya, 1 5 km W Perm (3 2, 2 3, WJP).
Ukrainian SSR: Crimea: Krymskiy Zapovednik ( 1 $, ZMMU).
Yugoslavia (4 2, 3 3): Croatia: Plitvice (RMNH), Skrad (GP).
Slovenia: Bled (RMNH), Cerknica (KS), Legatee (as Loitsch,
ZMH), Opcina Mtn. on Italian border E Triest (NHMV). Ser-
bia: Drazevac (FSAG).

Trypoxylon medium de Beaumont

Trypoxylon figulus var. medium DE BEAUMONT, 1945:477, 2,
3 (as media, incorrect original spelling). Holotype: 2, Swit-
zerland: Martigny (Mus. Zool. Lausanne). — BLUTHGEN, 1951:
234 (var. media); DE BEAUMONT, 1958:206 (forma media),
1959:30 (same); WOLF, 1959:15, 1 6 (figulus medium); VALK-
EILA, 1961:144 (var. media); DE BEAUMONT, 1964a:290,
1 964b:84 (forma media), 1965:56 (same), 1967:338 (same);
BOHART AND MsNKE, 1976:346 (ssp. medium); LOMHOLDT,
1976:267 (figulus media); RICHARDS, 1980:45 (var. media);
TSUNEKI, 1981:19 (medium).

Trypoxylon figulus minus var. rubi WOLF, 1959:15, 3. ! Lec-
totype, 3, Federal Germany: Plettenberg (coll. H. Wolf, Plet-
tenberg), present designation. Synonymized by DE BEAUMONT,
1964:291.

Trypoxylon figulus minus: WOLF, 1959:fig. c.

DIAGNOSIS.— There is no single character by
which medium can be distinguished from both
figulus and minus. The anteroventral mesotho-
racic carina is either straight or sinuate and curved
posterad mesally, but unlike that of most minus
it is not expanded into a process. Unlike those
of mosl figulus, the mesopleural setae around the
scrobe are shorter than the midocellar diameter.
In the female, the clypeal free margin is evenly
concave between the orbit and median projec-
tion (Fig. 7A), unlike that of minus and most
figulus, and the setae of the hindcoxal pit form
a curious channel-like structure visible only un-
der high magnifications (Fig. 1C, D) and which
has not been found in the other two species. The
hindcoxal pit is circular (Fig. 7B) instead of ob-
long (as it is in most figulus). The maximum
length of male flagellomere X equals 0.75-0.9 of
its basal diameter (the highest ratios are observed
in specimens in which the flagellomere XI is the
longest); the maximum length of flagellomere
XI is usually 2.0-2.2 times its basal diameter
(2.2-3.6 in figulus and minus). However, the fla-
gellomere XI length is 2.4 times its basal di-
ameter in some specimens, e.g., in a male from
Horsell, England (KMG), or a male from Uilac,
Romania; such specimens differ externally from
figulus in having a shorter mesopleural vestiture
and a longer flagellomere X. Gonoforceps with
ventroexternal expansion at about midlength (Fig.
7E, F); expansion absent in figulus and minus.

Body length 6.5-12 mm in female, 6.0-8.5 mm
in male.

LIFE HISTORY.— Several specimens of medium
(seen by me) were reared from Rubus twigs (Wolf,
1959). A male was reared from an old gall of the
chloropid fly Lipara lucens Meigen containing a
vacated nest of the bee Hylaeus pectoralis Forster
(England, Hampshire, Browndown, G. R. Else
collector, BMNH).

GEOGRAPHIC DISTRIBUTION (Fig. 8).— Western
and central Palearctic Region between Great
Britain and eastern Kazakh SSR, north to south-
ern England and beyond the Arctic Circle in Fin-
land, south to northern Mediterranean countries
including Mallorca, Crete, and Cyprus, as well
as Turkey, Syria, Israel, and northern Iran. Un-
known from Iberian Peninsula and North Africa.

RECORDS.— Austria (30 2, 20 3, NHMV if not indicated oth-
erwise): Burgenland: Andau (RMNH), Jois (JG), Donners-
kirchen an Neusiedlersee (KS), Neusiedl (KS, NHMV, ZMB),
Panzergraben an Neusiedlersee (JG), Winden (JG), Zurndorf
(AWE, RMNH). Niederosterreich: Bisamberg near Vienna,
Deutsch Altenburg (ZMB), Donauauen near Vienna (ZMH),
Hainburg (ZMB), Herzograd (JG), Marchfeld (JG), Oberwei-
den (DBB), Piesting, Schneeberg, Stammersdorf (DBB), Wien-
Tiirkenschanze. Oberosterreich: Gutau (JG). Salzburg: Kat-
chenberghohe (VH), Salzburg (BMNH). Tirol: Galtttr (BMNH),
Huben (ZMB), Innsbruck, Kraspes (DEI), 1 km SW Zwiesel-
stein. Voralberg: BielerhOhe.

Belgium (11 2, 173, FSAG): Antwerpen: Geel. Liege: Liege,
Montzen, Spa, Wandre. Limburg: Bree, Eben, Lummen. Lux-
embourg: Chatillon, Les Epioux, Torgny. Namur: Eprave, Fer-
age, Mont-Gauthier.

Bulgaria (1 2): Ruse (SMT).

Cyprus (3 2, 6 3): Amathus (FSAG, RMNH), Paphos (KMG).

Czechoslovakia (1 2, 2 3): Jihocesky Kraj: Prachatice (as
Prachatitz, NHMV). Vapadoslovensky Kraj: Sturovo (SMT).

Denmark (1 1 2, 8 3, ZMK, if not indicated otherwise): Born-
holm: Arnager. Frederiksborg: Hulerad, Humblebaek (S Hel-
singer), Tisvilde Hegn. Hjurring: Fauerholt Hede (ca 10 km
W Frederikshavn), Skoven on Lasse Island. Maribo: Kris-
tiansszede Skov on Lolland Island. Randers: Glatved Strand
and Kal0 on Djursland Peninsula, Mols Bjerge. Ribe: Bors-
mose, Ho Plantage. Svendborg: Keldsnoron Langeland Island.
Thisted: Bagsc at Nors (ca 7 km NW Thisted). Tender: Romo
Island (RMNH), Stensbaeck Plantage.

Finland (18 2, 15 3, HY if not indicated otherwise): Abo:
Pernio (AKM), Rymattyla (AKM). Alandia: Finstrom (AKM),
Hammarland. Karelia Australis: Virolahti. Karelia Borealis:
Nurmes. Kuusamo: Kuusamo (ZMK). Lapponia Inarensis: Iva-
lo. Nylandia: Hyvinkaa. Ostrobottnia Media: Paavola. Tavas-
tia Australis: Ha'meenlinna, Hattula, Janakkala, PSlkane, So-
mero (AKM), Vanaja.

France (24 2, 8 3, FSAG if not indicated otherwise): Aisne:
Liesse. Alpes Maritimes: Aurons (AWE). Aube: bois de Lignie-
res. Aveyron: Creissels. Basses-Alpes: Annot, Montagne de Lure
(ZMK), Peyresque. Dordogne: Paunat. Drome: Dieulefit
(RMNH). Haute-Loire: Pont de Sumene. Haut-Rhin: Rouf-
fach. Hautes-Alpes: Saint-Veran. Herault: Canet (BMNH).

PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS, MEDIUM, AND MINUS

137

FIGURE 7. Trypoxylon medium: A— female clypeus, B— female hindcoxa ventrally, C— female hindcoxal pit, vertical view,
D— same, oblique view, E— male genitalia (arrow: area shown in F), F— same, process of gonoforceps.

Landes: Mont-de-Marsan (MHNG), Parentis (KMG), St. Gi-
rons - Plage (RMNH). Moselle: Orny, Sierck. Pyrenees-Ori-
entales: La Llagonne - Mont Louis (VH). Var: Saint-Tropez,
Valescure (KMG). Vaucluse: Carpentras (RMNH).

Germany, Democratic (15 2, 6 <5): Berlin: Berlin (BMNH,
DEI, ZMB), Spandau (ZMB). Cottbus: Luckau (DEI). Dresden:
Gersdorf near Kamenz (SMT). Frankfurt: Buckow (ZMB),
Eberswalde area (DEI), Storitzsee near Berlin (DEI). Halle:
Bad Frankenhausen near Kyffhauser (HD), Halle (DEI), Kat-

tenburg near Kyffhauser (HD), Kyffhauser (DEI). Potsdam:
Gross Machnow (ZMB).

Germany, Federal (25 9, 34 <?): Baden-Wiirttemberg: Fed-
ersee (KS), Kaiserberg (ZMH), Kaiserstuhl (VH, ZMB), Karls-
ruhe (ZMH), Radolfzell (ZMH). Bayern: Balderschwang (KS),
Kreuth in Oberbayern (ZMB), Mainfranken (FIS), Miltenberg
(FIS), Obersdorf in Allgau (ZMB), Sondershausen (ZMH).
Hessen: Dillenburg (HW, WJP), Marburg (HW). Niedersach-
sen: Dorpen: 14 km SW Papenburg (VH), Norderney Island

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\~x4Bw

PULAWSKI: THE STATUS OF TRYPOXYLON FIGULUS. MEDIUM, AND MINUS

139

(VH). Nordrhein-Westfalen: Krefeld (DEI), Plettenberg (in-
cluding lectotype and 2 paralectotypes of rubi, HW, WJP),
Siegen (HW). Rheinland-Pfalz: Bad Munster am Stein (KS),
Mainz (KS).

Great Britain (53 5, 44 3, BMNH if not indicated otherwise):
Buckingham: Slough. Devon: Bovey Tracey, Braunton. Dorset:
Chideock, Wareham. Gloucester: Stroud (Rodborough).
Hampshire: Basingstoke. Bramdean, Browndown. Fareham,
Hum, Stockbridge. Isle of Wigt: Alum Bay, Sandown, St. He-
len's. Kent: Cobham, Darenth, Pluckley (KMG). London: Mill
Hill, Ruislip (KMG), Uxbridge. Oxford: Goring. Surrey:
Chobbham (KMG), Ewell (DBB), Horsell. Sussex: Findon,
Midhurst (Ambersham Common), Singleton (The Trundle).

Greece (1 9, 1 3): Crete: Kato Metokhi (ZMB). Peloponnesus:
Mega Spilaion (de Beaumont, 1965). Sterea Ellas: Kifissia
(KMG).

Hungary (2 9, 1 3): Pest: Ocsa (TMB). Tolna: Simontornya
(NHMV).

Iran ( 1 9): Gorgan, Shaskola Forest (JG).

Israel (3 9): Baniass (KMG), Eshtaol: Kesalon Valley (KMG).

Italy (199, 73): Emilia- Romagna: Modena: Zocca: Monte-
tortore (NHMV). Lazio (Frosinone): Pontecorvo (NHMV),
Roma (4 9, det. J. de Beaumont, WJP). Liguria: Genova
(MSNM), San Remo (GP). Lombardia (Varese): Mercallo
(MSNM). Piemonte: San Benedetto Belbo ca 20 km S Alba
(GP). Trentino-Alto Adige: Bolzano (as Bozen, NHMV), Ca-
valese (BB), Merano (as Meran, ZSM), Predazzo (ZMB). Ve-
neto: Colli Euganei: Montegrotto Terme (WJP), Garda (FSAG),
Jesolo (FSAG), Lido di Venezia (de Beaumont, 1959). Venezia
Giulia: Trieste (ZMH).

Netherlands (14 9, 103, RMHN if not indicated otherwise):
Drenthe: Emmen, Exloo, Mantinge. Gelderland: Ede (GVR),
Kootwijk, Hulshorst. Limburg: Heerlen. Noord Brabant:
Griendtsveen, Helenaveen, Udenhout, Waalwijk. Noord Hol-
land: Hilversum. Utrecht: den Dolder.

Norway (4 9, 5 3): Hordaland: Granvin (ZMUB). Sogd og
Fjordane: Balestrand (ZMUB). S«r-Tr»ndelag: Osen district
(FSAG).

Poland (1 3): Szczecin: Mi^dzyzdroje (as Misdroy, ZMB).

Romania (99, 18 3, MGA if not indicated otherwise): Brasov:
Uilac. Bucuresti: Valea Rosie near Oltenifa. Constanta: Ba-
badag (RMNH), Caraorman, Crisan, Gorgova in Danube Del-
ta (HD), Periprava, Valu lui Traian. Oradea: Oradea (RMNH).
Ploesti: Valea lui Bogdon near Sinaia.

Spain (1 9): Mallorca (DEI).

Sweden (22 9, 10 3, NRS): Asele Lappmark: Saxna's. Ble-
kinge: Ronneby. Bohuslan: Ljung. Dalarna: Falun. Gotland:
Fardume, Faro, Stanga. Halsingsland: jarvso. Jamtland: Stor-
lien. Oland: Hogby, Morbylanga, Vickleby. Skane: Ivo, Sten-
shuvud. Smaland: Kalmar Sund (Bla Jungfrun). Uppland: Bo-
gesund, Danderyd. Vastergotland: Osterplana hed.
Vastmanland: Skinnskatteberg.

Switzerland (20 9, 20 3, NHMB if not indicated otherwise):
Bern: Bern, Biel. Geneve: Bois d'Onex (MHNG), Genthod
(NHMB, RMNH), Peney (MHNG). Graubunden: Chur(ZSM),
Pare National Suisse (de Beaumont, 1958), Scanfs, Somvix.
Valais: Binntal (15 km NE Brig), Grimentz (RMNH), LOt-
schental, Martigny (BMNH), Saar Fee (BMNH), Sierre
(NHMV), Simplon (BMNH), Zermatt (RMNH). Vaud: Nyon
(MHNG).

Syria (1 3): Damascus (NHMV).

Turkey (4 9, 9 3): Amasya: Amasya (BMNH). Antakya: An-
takya (de Beaumont, 1967). Antalya: Antalya (BMNH), Finike

(de Beaumont, 1967), Finike-Ka? road (BMNH). Bursa: Ulu-
dag (BMNH). Kastamonu: Kastamonu area (BMNH). Mersin:
GOzne (BMNH), Mut (JG). Mugla: near KGycegiz (BMNH),
Marmaris (BMNH). Samsun: Samsun area (BMNH).

USSR: Armenian SSSR: Noyembryan (3 3, ZMMU). Geor-
gian SSR (2 2, ZMMU): Leselidze, Yermolovskoye near Gagra.
Kazakh SRR (79, 14 3, VLK if not indicated otherwise): Al-
maatinskaya oblast: 12 km W Alma Ata; Hi river 60 km E Hi
(now Kapchagai); Kargalinka, 5 km SW Alma Ata; Malaya
Almaatinka River in Zailiyskiy Alatau (ZMMU), 25 km S
Turgen village in Zailiyskiy Alatau. Semipalatinskaya oblast:
30 km SE Georgyevka; Tarbagatai Khrebet foothills 6 km N
Irinovka (which is 47°09'N, 81°53'E); 12 km N Zharma. Vos-
tochnokazakhstanskaya oblast: Baighym Canyon in Narym-
skiy Khrebet; Kalbinskiy Khrebet 20 km SW Leninka; Ken-
dyrlik river 1 5 km E Zaysan; 5 km N Oktyabrskiy in Ulbinskiy
Khrebet; 15 km NE Ust'-Kamenogorsk; 15 km SSW Ziry-
anovsk; 28 km SSE Ziryanovsk. Russian SSR: Bashkirskaya
Avtonomnaya oblast: Kazmash (1 3, ZMMU). Belgorodskaya
oblast: Valuyki ( 1 3, NHMV). Leningradskaya oblast: Primorsk
(as Koivisto, 1 2, AKM). Saratovskaya oblast: Kuznetsk (1 2,
ZMMU). Ukrainian SSR: Crimea (1 2, 2 3, ZMMU): Karadag,
Sebastopol.

Yugoslavia (6 9, 1 3): Croatia: Porec (FSAG), Puli (as Pola,
NHMV), Rab Island (as Arbe, TMB). Kosovo: Brezovica in
Sar Mts. (TMB), Pee (TMB). Serbia: Drazevac (FSAG), Sid
(FSAG). Slovenia: Portoroz (FIS).

LITERATURE CITED

BOHART, R. M., AND A. S. MENKE. 1976. Sphecid wasps of
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BLUTHGEN, P. 1951. Neues oder Wissenswertes iiber mittel-
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CRESSON, E. T. 1928. ThetypesofHymenopterain the Acad-
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DAY, M. 1979. The species of Hymenoptera described by
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DE BEAUMONT, J. 1945. Notes sur les Sphecidae (Hym.) de
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RICHARDS, O. W. 1980. Handbooks for identification of Brit-
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Ges. 18:11-16.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 11, pp. 141-158, 19 figs., 1 table January 17, 1984

DESCRIPTION AND OSTEOLOGY OF THRYSSOCYPRIS, A NEW

GENUS OF ANCHOVYLIKE CYPRINID FISHES, BASED

ON TWO NEW SPECIES FROM SOUTHEAST ASIA

Tyson R. Roberts

California Academy of Sciences, Golden Gate Park,
San Francisco, California 94118

and
Maurice Kottelat

Naturhistorisches Museum, Augustinerstrasse 2,
CH-4001, Basel, Switzerland

ABSTRACT: Thryssocypris new genus comprises two new species, the generic type-species '/'. smaragdinus
from the Kapuas River in Kalimantan Burnt (Indonesian western Borneo) and / . tonlesapensis from the lower
Mekong basin in Kampuchea and Vietnam. Extremely compressed and anchovylike, Thryssocypris differs
from all other cyprinids in having a non-protrusible upper jaw with the posterior half of its margin formed
solely by the maxilla, and a pair of elongate, slitlike supraethmoidal laterosensory trenches. Osteological
study reveals features of the Weberian apparatus, gill arches, pelvic girdle, and terminal radials of the dorsal
and anal fins that have not been reported previously in cyprinids. The two new species differ in proportions
and fin positions, in numbers of anal fin rays, scales, and vertebrae, and in coloration. The relationships of
the new genus to other cyprinids require further study.

INTRODUCTION Mekong species were obtained by F. d'Aubenton
Two recently discovered, anchovylike, insec- during a survey of the Tonle Sap in 1961, and
tivorous cyprinid fishes, one from the Kapuas at least one hundred specimens by W. J. Rain-
River in western Borneo (Kalimantan Barat, In- both during a University of Michigan fisheries
donesia) and the other from the lower Mekong survey in the Mekong Delta in 1974. This species
basin in Kampuchea and Vietnam, are closely also does not seem to have been collected pre-
related to each other and represent a highly spe- viously. The species differ so markedly in head
cialized new genus. We do not know of any pre- size and other characters that they were not im-
viously described species referable to this genus, mediately recognized as congeneric. However,
Five specimens of the Bornean species were closer comparison, including osteological study,
obtained at a single locality during an ichthyo- revealed that they agree in numerous characters
logical survey of the Kapuas River in 1976, and not found, or at least not reported, in any other
we have been unable to locate any additional cyprinids and that they clearly do belong in the
material of this species. Nine specimens of the same genus.

[141]

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 1

Cyprinidae is the largest family of freshwater
fishes, and we are very far from a phyletic clas-
sification of the genera or even a reasonable di-
vision of the family into taxonomic categories
above the generic level. A great deal of work
remains to be done in order to provide adequate
definitions for cyprinid genera, many of which,
as currently understood, are polyphyletic and
must be split up before a phyletic classification
can be achieved. Since many distinctive cyprinid
genera are relatively rare, and live specimens vir-
tually unobtainable, classification must be based
mainly on information that can be extracted from
preserved specimens. Osteological accounts of
distinctive genera, especially if extensively illus-
trated in a standardized format (e.g., that of Bry-
con by Weitzman 1 962) provide perhaps the most
ready source of information for use in phyletic
classification. Unfortunately, there is a dearth of
such accounts. In addition to describing this new
genus and its two species, therefore, we present
an account of its osteology.

MATERIAL AND METHODS

Material of the two species described herein is
deposited in the following institutions: Califor-
nia Academy of Sciences (CAS), Museum of Zo-
ology of the University of Michigan (UMMZ),
Smithsonian Institution (USNM), Museum na-
tional d'Histoire naturelle, Paris (MNHN), Mu-
seum d'Histoire naturelle de Geneve (MHNG),
Museum Zoologicum Bogorense, Bogor, Indo-
nesia (MZB). The osteological account is based
on a 53.2-mm paratype of T. smaragdinus and
a 46.0-mm paratype of T. tonlesapensis, which
were cleared and stained in alcian blue and aliza-
rin. Additional osteological observations were
made on radiographs of the other four type spec-
imens of T. smaragdinus and 1 2 of the largest
T. tonlesapensis. Although our osteological fig-
ures are based mainly on T. smaragdinus, our
remarks generally relate to the osteology of both
species. While it is not always so in fishes pre-
pared with alcian and alizarin, in our material
of T. smaragdinus and T. tonlesapensis, except
for the distal ends of the intermuscular bones, it
seems that all skeletal elements stained with al-
cian are true cartilage; all stained with alizarin
are true bone. In the figures bone is indicated by
stipple, cartilage by simple diagonal hatching,
and fenestrae or foramina opening into intracra-

nial spaces by cross-hatching. In all of the figures
the scale bar equals 1 mm.

We draw attention to a few reservations con-
cerning the osteology. The parietal laterosensory
canal bony tubules and supraoccipital crest were
badly damaged in the 53.2-mm specimen illus-
trated and were drawn on the basis of reconstruc-
tion and comparison with whole, unstained spec-
imens. The shape of the posteriormost portions
of the prevomer and parasphenoid and their re-
lationships to other elements were not observed
with certainty in the specimen illustrated and
may be in error. In particular, the appearance of
the prevomer may be due to breakage rather than
to mere separation. We did not detect an inter-
calar, an element sometimes absent in Cyprini-
dae, but are uncertain of its absence in the present
instance. Otoliths were not removed so that the
crania could be preserved intact, and thus they
are not described or figured.

Thryssocypris, new genus

TYPE-SPECIES. — Thryssocypris smaragdinus Roberts and Kot-
telat, new species.

DIAGNOSIS. — Small (largest specimen 63.8
mm), anchovylike cyprinids with highly com-
pressed head and body; barbels absent; excep-
tionally large olfactory organ; well-developed
hyaline eyelid; elongate, moderately upturned,
and very narrow terminal jaws; acutely pointed
snout, slightly to strongly projecting anterior to
upper jaw; low dorsal and anal fins, with falcate
margins, originating in posterior half of body;
abdomen rounded, without keel. Thryssocypris
differ from all other known cyprinid genera in
the following characters: 1) upper jaw entirely
nonprotrusible, without rostral cap or even ves-
tigial groove of rostral cap, with posterior half
of border formed solely by maxilla; 2) cephalic
laterosensory system with elongate, slitlike su-
praethmoidal or rostral canal medial to nasal
canal, supraorbital canals similarly slitlike, not
enclosed in bony tubules on frontal bone, and
an elongate dermosphenotic canal enclosed in a
bony tubule fused for entire length to dorsal sur-
face of sphenotic bone; 3) ventral portion of gill
arches highly specialized, ceratobranchials 1-4
with proximal portions abruptly narrowed and
ending in elongate cartilaginous extensions, hy-
pobranchial one minute and hypobranchials 2-3

ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS

143

FIGURE 1. Thryssocypris smaragdinus, 49.4-mm holotype (MZB 3435).

absent, and basibranchials extremely slender;
4) Weberian apparatus with lateral process of
centrum 2 extremely elongate and strongly curved
posteriorly dorsal to tripus, extending laterally
equally as far as fully formed pleural rib of fifth
vertebra; 5) ischiac process of pelvic girdle with
elongate and extremely slender anterior and pos-
terior processes; and 6) posteriormost pterygio-
phore of dorsal and anal fins with a deeply bi-
furcate radial projecting considerably beyond base
of posteriormost fin rays.

Regarding the diagnostic characters listed
above: 1 ) a few other cyprinids have nonprotru-
sible upper jaws (e.g., the North American Exo-
glossum and Parexoglossum), but in these and
all other cyprinids known to us the maxilla is
entirely or almost entirely excluded from the gape
by the premaxilla; 2) some other cyprinids ap-
parently have laterosensory canals in the su-
praethmoidal region (e.g., Luciosomd), but the
position and shape of these canals is quite dif-
ferent from those in Thryssocypris, and they are
not slitlike. Lekander ( 1 949) reported rostral lat-
erosensory organs in some European cyprinids,
but these are more anterior (near snout tip) and
are joined by a commissure (absent in Thrys-

socypris). Supraorbital canals in Cyprinidae are
usually enclosed in bony tubules on the dorsal
surface of the frontal bone. A dermosphenotic
laterosensory canal is present in most cyprinids,
but usually occurs in a short segment of bony
tubule or on a small laminar dermosphenotic
bone that is completely separate and superficial
to the sphenotic bone; 3) in all other cyprinids
we have examined or know about the gill arches
have ceratobranchials 1-4 uniformly wide and
hypobranchials one to three present; 4) many
cyprinids have a very large lateral process on
centrum two, but in most instances it projects
directly laterally from the vertebral column, or
in some instances is strongly curved posteriorly
but projects ventrally to the tripus (rather than
dorsally to it as in Thryssocypris}; 5-6) the un-
usual condition of the ischiac process and of the
terminal radials in the dorsal and anal fins is
unlike anything we know of or have seen reported
in any other cyprinids.

A more detailed account of the osteology of
Thryssocypris is given following the species de-
scriptions. Some additional characters of the ge-
nus are given here. Lateral line complete, mod-
erately curved downward anteriorly. Gill rakers

FIGURE 2. Thryssocypris tonlesapensis, 50.6-mm holotype (MNHN 1982-1032).

144

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 1
TABLE 1 . QUANTITATIVE CHARACTERS IN Thryssocypris.

Character

T. smaragdinus

T. tonlesapensis

Counts

Gill rakers on first gill arch1
Pharyngeal teeth (left/right)'
Dorsal fin rays
Anal fin rays
Pectoral fin rays
Pelvic fin rays
Procurrent caudal fin rays
Scales in lateral series
Scale rows above + below lateral line
Median predorsal scales
Circumferential scales
Circumpeduncular scales
Abdominal + caudal = total vertebrae

0 + 7
1+5/5+1

0+8

2 + 5/5 + 2
9'/2(3),10'/2(9)

11-12

7-8

9 + 9

36-39

5 + 3

257-27?

18-20

14-15

21 + 17 = 38(2),21 + 18 = 39(3)

10-12

7-8

9 + 8-9
43-46

6 + 3
26-31
18-21
16-17
21+22 = 43(12),21+23 = 44(1)

Proportions (times in standard length)

Head

3.8-4.1

4.9-5.3

Snout

12.0-12.5

15.4-16.9

Olfactory organ

18.4-22.4

33.6-34.7

Eye

15.5-17.0

19.2-24.8

Lower jaw'

7.9

10.7

Pharyngeal bone1

11.4

20.0

Body depth

5.1-5.5

6.2-6.8

Body width

9.9-11.1

12.2-14.3

Caudal peduncle depth

11.4-12.0

10.6-11.5

Pectoral fin

5.7-6.1

6.0-7.0

Pelvic fin

9.0-10.0

8.9-9.7

Preanal length

1.3-1.4

1.5-1.6

Other

Inclination of jaws

25°

30-35°

1 From cleared and stained specimens.

small, short, and somewhat stubby, leading edge
of lower limb of first gill arch with seven to eight
rakers, upper limb with none to one. Pharyngeal
teeth uncinate, in two rows, 1 -2 + 5/5 +2-1 . Scales
large, 36-46 in lateral series, approximately cor-
related in number with vertebrae, which total
38-44. Quantitative characters of the genus are
summarized in Table 1 .

Dorsal and ventral profiles of head and body
anterior to dorsal and anal fins uniformly and
gently curved (not forming an angle at occiput
or pectoral fin origin). Dorsal and ventral surface
of body anterior to median fins rounded from
side to side. Body moderately tapered caudally
(markedly tapered in some cheline cyprinids).
No indication of a cranial flexure. Morphological
features related to ability of head to tilt upwards
in relation to vertebral column (evidently a feed-
ing adaptation in Asian cyprinids such as Mac-
rochirichthys, Salmostoma, Oxygaster, and Che-

Id) are absent. Epaxial musculature does not
invade cranial roof, angle of occiput is relatively
acute, and parasphenoid relatively horizontal.

Scales on dorsum, sides (including lateral line
scale series), caudal peduncle, and abdomen
morphologically similar, except that scales of lat-
eral line series have simple tubule for laterosen-
sory canal, horizontally oriented, originating at
or near focus and extending for half or more than
half length of posterior field. Shape modified oval;
anterior, dorsal, and ventral margins slightly
convex, posterior margin rounded or even slight-
ly pointed (especially in median scale rows and
near bases of paired and median fins), sometimes
weakly scalloped. Dorsal and ventral fields more
or less sharply set off from anterior field but grad-
ing smoothly into posterior field. Radii present
on all fields, but best developed on anterior and
posterior fields. Radii on anterior field 8-15, hor-
izontally oriented (parallel to each other). Radii

ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS

145

-••„...

»rV

" '.'"'•- •-/.,]
\

FIGURE 3.

optics).

Thryssocypris smaragdinus, 49.7 mm, twenty-third scale in lateral line scale row (Nomarski interference contrast

on posterior field 22-30 or more, slightly diver-
gent from horizontal, more widely separated than
those on anterior field. Radii on anterior and
posterior fields originating near focus. Radii on
dorsal and ventral fields similar, few in number,
widely divergent, and variable in length, origi-
nating remote from focus, nearly parallel to up-
permost and lowermost radii in posterior field,
with which they seem to form a continuous se-
ries, and entirely divergent from radii in anterior
field. Circuli well defined and evenly spaced in
anterior, dorsal, and ventral fields, but discon-
tinued or indistinct on posterior field. Circuli of
anterior field vertical, of dorsal and ventral fields
horizontal; circuli of dorsal and ventral fields
meeting at right angles with circuli of anterior
field at interfield margins, and bisecting growth

lines of posterior field at a sharp angle. Growth
lines, readily observable in posterior field, much
more numerous and more nearly circular in ar-
rangement than circuli.

Multicellular horny projections, or tubercles
(also known as breeding tubercles), absent from
body and fins, and perhaps also from head. Nu-
merous minute conical projections toward tip of
snout, on upper lip, and on lacrimal area of head
(especially near ventrolateral margin of infraor-
bital 1) appear to be tubercles. Tubercles fre-
quently occur on dorsal surface of pectoral fins,
on scales, and on mandible in many cyprinids,
especially in males, but are absent from these
places in Thryssocypris.

ETYMOLOGY.— From the Greek thrissos, a her-
ring or anchovy, and cypris, a small minnow.

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FIGURE 4. Thryssocypris, radiographs, a, T. smaragdinus, 49.4 mm (holotype); b, T. tonlesapensis, 52.0 mm (paratype, CAS
50946). Note obvious differences between species but also basically identical morphology of skulls.

Thryssocypris smaragdinus, new species

(Figures 1, 3a, 4-9, lOb, 11-19)

HOLOTYPE. — MZB 3435, 49.4 mm, mainstream of upper
Kapuas River, 6 km w of Putussibau, Kalimantan Barat, In-
donesia, lat. 0°50.5'N, long. 1 12°52'E. Seining in late afternoon
and at dusk, on gently sloping open beach with coarse sand to
fine gravel bottom and moderate current. 9 Aug. 1976.

PARATYPES.-CAS 49314, 3: 46.8-54.0 mm, and USNM
230243, 1: 49.7 mm, collected with holotype.

DIAGNOSIS.— Thryssocypris smaragdinus is
most readily distinguished from T. tonlesapen-
sis, its only congener, by much larger head, larger
eye, larger pharyngeal bone, and fewer anal fin
rays, scales, and vertebrae. These and other
quantitative characters differing in the two species
are summarized in Table 1 . In T. smaragdinus,
anal fin origin on a vertical with dorsal fin origin
(versus anal fin origin far in advance of dorsal
fin origin in T. tonlesapensis). Snout tip projects
strongly beyond upper jaw in four of the five
specimens of T. smaragdinus (including the ho-
lotype), thus differing strikingly from T. tonle-
sapensis, in which it projects but very slightly.
In one paratype of T. smaragdinus (USNM
230243, 49.7 mm), however, the snout tip pro-
jects little more than in T. tonlesapensis. Some
additional differences between the two species
are indicated in the color descriptions below and
in the osteological account following.

In life T. smaragdinus are brilliant emerald
green on the upper half of the head and body
and bright silvery below. Preserved specimens
exhibit a wide longitudinal band, narrowest an-

teriorly, extending from head to caudal fin, and
lying entirely in dorsal half of body. Middle of
caudal peduncle and caudal fin base with dark
round spot level with longitudinal band. Mela-
nophores absent from all fins except for a few
small scattered ones on interradial membranes
of dorsal and caudal fins. Melanophores almost
entirely absent on ventral half of body. Dorsal
half of body with numerous fine melanophores
or chromatophores in addition to those of lon-
gitudinal band, but not forming noticeable pat-
terns such as rows parallel to posterior margin
of scales. Dorsal midline of body with two or
three thin longitudinal rows of melanophores.
Dorsal surface of head, especially dorsal to nasal
organs, deeply pigmented with numerous large
melanophores. Inner surface of opercle dusky.

Gut contents of 53.2-mm paratype comprise
moderately numerous triturated remains of
aquatic coleopteran and dipteran larvae, and
some adult winged dipterans. No other food items
observed.

ETYMOLOGY.— From the Greek smaragdinos,
emerald green.

Thryssocypris tonlesapensis, new species

(Figures 2, 3b, lOa)

HOLOTYPE.— MNHN 1982-1032, 50.6 mm, Prek Tamen, at
or near Snoc Trou, Kampuchea, 9 Nov. 1961.

PARATYPES.-MNHN 1982-1033, 3: 44.7-50.3 mm, MHNG
21 19.63-64, 2: 45.5-46.0 mm, CAS 50946, 2: 48.4-52.1 mm,
collected with holotype; MNHN 1982-1034, 1: 52.3 mm, Prek
Tasom, at or near Snoc Trou, Kampuchea, June 1961; UMMZ

ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS

147

dermosphenotic

extrascapular

posttemporal

supracleithrum

nasal

frontal

ethmoid

basioccipital
exoccipital

lateral ethmoid

sphenotic parietal pterotic
FIGURE 5. Thryssocypris smaragdinus, 53.2 mm, cranium (dorsal view).

epiotic

210277, 1: 43.4 mm, Mekong River at w end of Con Phong
(or Con Ho?) Island 2-5 km w of My Tho, Vietnam, 18 July
1974; UMMZ 210278, 1: 45.7 mm, Mekong River at Con Ho
Island, My Tho, Vietnam, 19 June 1974; UMMZ 210279, 58:
20.4-60.1 mm, fish market at Vinh Long, Vietnam, 22 June
1974; UMMZ 210280, 36: 29.2-63.8 mm, fish market at Can
Tho, Phong Dinh Province, Vietnam, 23 June 1974.

DIAGNOSIS. — Characters distinguishing T.
tonlesapensis from T. smaragdinus are given in
the diagnosis of T. smaragdinus, in Table 1, in
the description of coloration and other remarks
below, and in the osteological account following.
The species is immediately distinguished from
T. smaragdinus by its much smaller head, more
numerous scales (44-46 in lateral line series ver-
sus 36-39), and more numerous anal fin rays
(\5l/2-\6l/2 versus 12'/2-14'/2). The very striking
difference in size of the pharyngeal jaws of ton-
lesapensis (Fig. 4), the linear dimensions of which
are only about half as great as in T. smaragdinus,
is much greater than would be expected if this
difference were due only to the difference in head
size, since the head is only about 20% longer in
T. smaragdinus.

Coloration of live T. tonlesapensis has not been
observed. Specimens observed in fish markets
by W. J. Rainboth were entirely silvery. Color
pattern of preserved specimens is similar in its
basic features to that of T. smaragdinus, includ-
ing the longitudinal band and basicaudal spot,
but differing in some details: longitudinal band
lower, nearly midlateral in position (confined to
dorsal half of body in T. smaragdinus); mela-

nophores on dorsal half of body tending to form
rows parallel to posterior margins of scales; and
inside of opercle clear instead of dusky.

Guts of numerous specimens are moderately
to very full of insects, mostly aquatic larvae; no
other food items observed.

ETYMOLOGY.— From Tonle Sap, the enormous
permanent backwater of the lower Mekong, into
which the Prek Tamen and Prek Tasom flow.

OSTEOLOGY

Although the two species of Thryssocypris dif-
fer strikingly in skull size and there are obvious
differences in the axial skeleton and median fin
skeletons related to differences in vertebral num-
ber and anal fin position and ray number, their
osteology is very similar in most respects (Fig.
4). Some osteological differences between the two
species are noted in the following account, which
is based on both species, even though the draw-
ings (except Fig. lOa) are of T. smaragdinus.

CRANIUM (Figs. 5-8, 10). — Roof of cranium
entire, without fontanel, strongly convex trans-
versely for its entire length. Frontals with nar-
row, flangelike, nearly horizontal lateral margins,
and a transverse shallow depression or groove
overlying tectum cranii or epiphyseal bar. Sphe-
notic as well as pterotic bones contribute sub-
stantially to cranial roof, with moderately de-
veloped sphenotic and pterotic projections or
spines. Ethmoid (or supraethmoid) very large,

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 1

frontal sphenotic

orbitosphenoid

ethmoid
\
preethmoid

prevomer

mesethmoid

dermosphenotic

parietal

supraoccipital

epiotic

pterotic

lateral ethmoid

pterosphenoid

parasphenoid

1 basioccipital

FIGURE 6. Thryssocypris smaragdinus, 53.2 mm, cranium (lateral view).

completely covering mesethmoid so that it does
not contribute to dorsal surface of ethmoid re-
gion. Preethmoids cartilaginous. Dilatator fossae
indistinct. Posttemporal fossae absent.

In most cyprinids the cranium tends to be dor-
sally flattened, or even transversely concave. In
rasborines and bariliines the cranium tends to
be barrel-shaped (Gosline 1975), with a convex
dorsum, as in Thryssocypris. The sphenotic is
usually entirely or almost entirely excluded from

the dorsal roof of the cranium, a noted exception
occurring in Esomus (Ramaswami 1955), which
in other respects differs very much from Thrys-
socypris. In Cyprinidae the ethmoid is usually
much shorter than in Thryssocypris, broader than
long, and with its anterior margin broadly and
deeply indented medially to receive the kineth-
moid (Thryssocypris has only a very small in-
dentation anteriorly; see Fig. 5).
In chelines (Howes 1979) the cranium is more

ethmoid
prevomer

optic foramen

orbitosphenoid
i

lateral ethmoid

pterosphenoid
sphenotic

exoccipital
basioccipital

preethmoid

\
mesethmoid \ /

nasal frontal

lUOOd

Thryssocypris smaragdinus, 53.2 mm, cranium (ventral view).

parasphenoid hyomandibu|ar
fossa

FIGURE 7.

ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS

149

parietal

supraoccipital

epiotic
pterotic

exoccipital
basioccipital

lateral occipital
fenestra

foramen magnum

cavum sinus imparis
aortic canal

FIGURE 8. Thryssocypris smaragdinus, 53.2 mm, cranium (occipital view).

or less flat or even concave dorsally, the mes-
ethmoid forms a shelflike projection extending
anteriorly to the ethmoid, the anterior half of the
parasphenoid is oriented at an angle of about 20°
to 30° from the horizontal, the supraoccipital
crest tends to be dorsal in position, and the oc-
ciput is elongated and gently sloping. In all of
these respects chelines differ from Thryssocypris.
JAWS (Figs. 9-11).— Jaws elongate, upturned,

and very narrow. Premaxilla only half as long as
maxilla, maxilla alone forming margin of pos-
terior half of upper jaw (maxilla entirely excluded
or almost entirely excluded from gape in all other
cyprinids). In T. smaragdinus maxilla with two
ascending or ethmoid processes, each with a car-
tilaginous cap, anterior process moderately elon-
gate or pedicellate, posterior process short (Fig.
1 1); in T. tonlesapensis maxilla with only a single

infraorbitals 1-5

supraorbital

nasal

suprapreopercle
opercle

premaxilla

maxilla

dentary

subopercle
interopercle

FIGURE 9. Thryssocypris smaragdinus, 53.2 mm, jaws and facial bones (lateral view).

quadrate

preopercle

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 1

premaxilla
palatine

maxilla

infraorbital 1
lateral ethmoid

nasal

kinethmoid
prevomer
preethmoid

ethmoid
frontal

FIGURE 10. a, Thryssocypris tonlesapensis, 46.0 mm, ethmoid region of skull (dorsal view); b, Thryssocypris smaragdinus,
53.2 mm, kinethmoid bone (dorsal view).

ethmoid process, short and broad based (non-
pedicellate), and with a cartilaginous cap (Fig.
10). Anterior half of maxilla with a medially di-
rected shelf arising from its dorsal margin, coex-
tensive with and largely overlapping lateral sur-
face of premaxilla. Maxilla with a slender,
posteriorly curved, broad-based projection aris-
ing near middle of its dorsal surface and extend-
ing medially to infraorbital number one, to which
it is firmly attached by connective tissue, thus
contributing to immobilization of upper jaw. Only
posterior half of maxilla free from snout and
capable of limited movement. Posterior portion
of maxilla, separated from infraorbitals by a shal-
low groove, simple, elongate, and flexible; ex-

tends to proximal end of dentary. Lower jaw
slightly shorter than upper jaw, elongate and very
narrow, completely included within gape when
mouth is closed. Symphysis of lower jaw without
pronounced dorsal or ventral knobs, although
vestigial or rudimentary dorsal symphyseal knob
may be present, especially in T. tonlesapensis.
Dorsal margin of dentary with a slight elevation
just behind symphysis in T. tonlesapensis; in T.
smaragdinus elevation absent or barely notice-
able. Dentary and angular with coronoid pro-
cesses short, vertically oriented, and separate,
that of dentary arising very near proximal end
(coronoid process of dentary frequently very large,
usually arising near middle of dentary, and not

, premaxilla

maxilla

angular

dentary

Meckel's cartilage

, , coronomeckelian retroarticular

FIGURE 1 1. Thryssocypris smaragdinus, 53.2 mm, jaws (medial view).

ROBERTS & KOTTELAT: THRYSSOCYPRIS. A NEW CYPRINID GENUS

151

preopercle
FIGURE 12. Thryssocypris smaragdinus, 53.2 mm, suspensorium (lateral view).

separate from angular in most cyprinids; angular
without distinct coronoid process in most cyp-
rinids). Free ventromedian margins of dentary
nearly straight for entire length, those of opposite
sides in contact or closely approximated (ven-
tromedian margin of dentary frequently highly
modified in shape in cyprinids, as in Danio and
its relatives, and dentaries of opposite sides usu-
ally more or less widely separated in cyprinids,
including genera with comparably elongate and
narrow lower jaws such as Macrochirichthys,
Salmostoma, and Securicula).

The poorly developed condition of the single
ethmoid process on the maxilla of T. tonlesa-
pensis is suggestive of the condition in chelines,
but because the jaws of Thryssocypris are oth-
erwise very unlike those of chelines and because
T. smaragdinus has well-developed ethmoid
processes, this resemblance is probably due to
independent reduction (possibly related to small
size of the head as well as nonprotrusibility of
the jaws).

Development of large dorsal symphyseal knobs
on each dentary (and sometimes almost equally
prominent ventral symphyseal knobs) is char-
acteristic of most chelines, and small dorsal sym-
physeal knobs occur in many cyprinids, includ-
ing Rasbora and Barilius. In forms with dorsal
symphyseal knobs the symphysis of the upper
jaw is frequently indented for their reception;

there is no comparable indentation in the upper
jaw of Thryssocypris.

INFRAORBITAL AND NASAL BONES (Fig. 9).—
Infraorbital series complete, with five large, la-
mellar infraorbital bones each bearing a simple
bony tubule for infraorbital laterosensory canal
and, together with supraorbital bone, forming an
almost complete ring around eye. Posterior bor-
der of supraorbital indented where concave an-
terior margin of infraorbital 5 fits snugly against
it; slender, pointed, anteroventral projection of
supraorbital nearly contacts dorsal margin of in-
fraorbital 1 . Nasal bone moderately elongate, with
a narrow lamellar portion on either side of bony
tubule for nasal laterosensory canal.

Among Asian minnows with elongate, up-
turned jaws and compressed bodies, a complete
infraorbital series in which all five infraorbitals
have well-developed lamina is typical of rasbo-
rines and bariliines and unusual in chelines (Gos-
line 1975). In Cyprinidae the primitive number
of infraorbitals, exclusive of the dermosphenotic
(sometimes counted as an infraorbital), is five.
Occasionally a greater number occurs, but this
is usually (perhaps invariably) due to fragmen-
tation of more or less tubular infraorbitals in
which the lamellar component is greatly reduced
or absent.

SUSPENSORIUM (Fig. 12). — Palatine arch nearly
horizontal. Hyomandibular and preopercle ver-

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urohyal

dorsal hypohyal
ceratohyal

ventral hypohyal

branchiostegal rays 1-3
FIGURE 13. Thryssocypris smaragdinus, 53.2 mm, hyoid bar and urohyal (lateral view).

tical. Lateral abductor process of hyomandibular
dorsally directed, originating just below hyo-
mandibular articulation with cranium. Meta-
pterygoid-quadrate fenestra absent or greatly re-
duced and almost entirely occluded by peculiarly
expanded distal end of symplectic. Anteroven-
tral limb of preopercle extremely short. Quadrate
nearly square, with strong posteroventral exten-
sion seen in many or most cyprinids, and without
a foramen immediately posterior to articular fac-
et for lower jaw. Ectopterygoid apparently absent
in T. smaragdinus; present but reduced in T.
tonlesapensis. Palatine bone with a well-devel-
oped posteroventral extension underlying lateral
ethmoid articular facet. An elongate, continuous
palatine cartilage connects palatine bone to me-
sopterygoid anteriorly and to metapterygoid and
quadrate posteriorly. At its anterior end, between
palatine bone and mesopterygoid, this cartilage
forms a large, saddlelike facet articulating with
ventral surface of lateral ethmoid. Opercle gen-
eralized in shape, deeper than long, its posterior
margin rounded, dilatator process weakly de-
veloped. Suprapreopercle short, tubular, fused to
anterodorsal corner of opercle.

In chelines the palatine bar is usually upturned
like the strongly upturned jaws, and is thus
obliquely oriented. Many cyprinids, including
some chelines and bariliines, have a well-devel-
oped metapterygoid-quadrate fenestra, and most
have the quadrate with a well-developed pos-
teroventral process, a preopercle with a promi-
nent anteroventral portion, and a large, broad

ectopterygoid. In T. smaragdinus the elongate
ventral portion of the mesopterygoid extends
narrowly below the palatine cartilage where the
ectopterygoid normally occurs, but examination
with direct and transmitted light failed to reveal
an ectopterygoid. In T. tonlesapensis the meso-
pterygoid does not extend ventral to the palatine
cartilage, and a very thin, elongate ectopterygoid
is present. Most chelines (Howes 1979) and Lu-
ciobrama (Howes 1978) have a foramen in the
quadrate immediately posterior to its articula-
tion with the lower jaw. Chelines and some other
cyprinids tend to have a stout, fingerlike dilatator
process forming the elevated anterodorsal corner
of the opercle. Some cyprinids have a free su-
prapreopercle and some lack this element alto-
gether. Fusion of the Suprapreopercle with the
opercle occurs frequently but is known only in
the Cyprinidae. The cyprinid Suprapreopercle is
always a simple bony tube enclosing the upper-
most portion of the preopercular laterosensory
canal, never with a laminar portion (sometimes
greatly enlarged) as in characoids.

HYOID BAR AND UROHYAL (Fig. 13). — Hyoid
bar generalized for Cyprinidae, with three bran-
chiostegal rays as in all members of the family,
and no unusual features. First branchiostegal ray
articulated to ventral surface of ceratohyal, which
has a notch in margin at point of articulation;
second branchiostegal ray broadly articulated to
lateral face of ceratohyal; third broadly articu-
lated to lateral face of epihyal. Basihyal dorso-
ventrally compressed, its anterior third cartila-

ROBERTS & KOTTELAT: THRYSSOCYPRIS. A NEW CYPRINID GENUS

153

hypobranchial 1
basibranchials1-2

infrapharyngobranchials 2-3
ceratobranchials 1-5

basihyal

epibranchials 1-4

FIGURE 14. Thryssocypris smaragdinus, 53.2 mm, gill arches and pharyngeal jaws (dorsal view, with upper half of arches
on left side deflected to reveal ventral surface).

ginous and with a broadly rounded anterior
margin (basihyal elongate and narrow for its en-
tire length in at least some chelines). Interhyal
tubular and moderately elongate (flattened and
laterally expanded in some chelines and cul-
trines). Urohyal with posteroventral process not
greatly expanded, its distal end simple in T.
smaragdinus and with a very small notch in T.
tonlesapensis. (Urohyal greatly modified in some
bottom dwelling cyprinoids including cyprinids;
posteroventral process very deeply forked in Lu-
ciobrama and at least some chelines.)

GILL ARCHES AND PHARYNGEAL JAWS (Fig.
14).— Gill arches (main features described above
under generic diagnosis) basically similar in T.
smaragdinus and T tonlesapensis. Narrow prox-
imal portion of ceratobranchials not quite so long
as in T. smaragdinus, but equally strongly nar-
rowed and with peculiar cartilaginous proximal
ends as large as in T. smaragdinus. Basibran-
chials two in T. smaragdinus, three in T. ton-

lesapensis, extremely slender in both species.
Ceratobranchial 5 (tooth-bearing pharyngeal
bone) uniformly slender, its length more than
five times its width, gracefully arched. External
ala elongate but extremely narrow, its origin
marked by a small, strongly angular projection
opposite base of middle tooth in major tooth
row. Ventral edentulous limb slightly shorter, and
dorsal edentulous limb slightly longer, than den-
tigerous portion. Symphyseal half of ventral
edentulous limb abruptly narrowed and set at
angle to rest of bone, so its orientation is nearly
horizontal. Pharyngeal teeth uncinate (conical
with recurved tips), in two rows; those of inner
or major row uniformly decreasing in size from
front to back, those of minor or outer row sub-
stantially smaller than those of major row.

Like most cyprinids Thryssocypris has but two
infrapharyngobranchials, here interpreted as in-
frapharyngobranchials 2-3. They exhibit the
characteristic two-plus-two relationship with

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np2-3

neural
complex

scaphium

intercalarium

supraneural

prezygapophysis

Ipcl
Ipc2

basioccipital tripus

FIGURE 15. Thryssocypris smaragdinus, 53.2 mm. Weberian apparatus (lateral view) (Ipcl. 2 = lateral processes of centrum
1 and centrum 2: np2.3 = neural plates of centrum 2 and centrum 3).

os suspensorium

epibranchials 1-2 and 3-4 which seems to be
characteristic of all cyprinoids. Epibranchials 1-
2 are united by cartilaginous joints to the distal
end of infrapharyngobranchial 2, and epibran-
chials 3-4 are similarly joined to infrapharyn-
gobranchial 3. Thus the proximal ends of infra-
pharyngobranchials 1 and 2 approximate each
other, and there is a slight but distinct gap be-
tween epibranchials 1-2 and epibranchials 3-4.
This peculiar orientation, which seems not to
have been noted previously, has been observed
without exception in numerous cyprinoids ex-
amined by us. Although the orientation is highly
specialized compared to the primitive one-to-
one relationship between the infrapharyngobran-
chials and epibranchials observed in characoids
and most lower teleosts, it is apparently primi-
tive for cyprinoids. A curiously similar two-plus-
two arrangement is present in numerous silu-
roids examined, including Diplomystes, but the
relationships of the infrapharyngobranchials to
the epibranchials appears not to be exactly the
same. Further investigation should be done to
determine whether this condition in cyprinoids
and siluroids indicates a common ancestor for
these two groups. In any event this specialized
condition is too widespread among cyprinids to
be of any use in assessing phyletic relationships
of Thryssocypris.
In some cyprinoids (mainly bottom feeders,

such as Labeo and Osteochilus, with inferior,
suctorial mouths) the basibranchial bones appear
to be shaped and articulated in such a fashion as
to permit them to slide over and under each
other, thus indicating that the ventral half of the
branchial basket is capable of anteroposterior
contraction and expansion. In Thryssocypris the
elongate, slender, and straight basibranchial bones
are firmly joined end to end, indicating that the
branchial basket is incapable of such contraction
and expansion movements.

WEBERIAN APPARATUS (Fig. 1 5). — Similar gen-
erally to Weberian apparatus of Opsariichthys
(Fink and Fink 1981; Fig. 14). Neural complex
vertical (not strongly posteriorly sloped as in
many chelines and cultrines), widely separated
from occiput. Occiput attached to anterodorsal
surface of Weberian apparatus by a thin median
strip of cartilage extending from posterodorsal
median margins of exoccipitals to claustrum and
neural process of centrum number two. Lateral
process of centrum 2 exceptionally elongate and
strongly curved posteriorly, extending dorsally
to tripus and laterally as far as major curvature
of fully formed pleural rib of vertebra 5. Fourth
pleural rib and parapophysis (bearing os suspen-
sorium) without a prominent anterodorsally pro-
jecting lateral process (present in Opsariichthys).
Intercalarium simple, slender, and elongate, hor-
izontally oriented, without ascending or articular

ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS

155

post temporal

extrascapular

supracleithrum

mesocoracoid
postcleithrum

scapula

coracoid

FIGURE 16. Thryssocypris smaragdinus, 53.2 mm. pectoral
girdle, left half (lateral view).

processes. Centra 2 and 3 fused to each other, at
least ventrally (centra 2-3 separate in Opsari-
ichthys but partially or completely fused in many
cyprinids).

Endochondral union of Weberian apparatus
with cranium is characteristic of cyprinoids and
is not found in other ostariophysans (Fink and
Fink 1981). Reduction and loss of ascending and
articular processes of intercalarium in other os-
tariophysans, especially siluroids, is reported by
Fink and Fink (1981:328). In Opsariichthys rel-
atively large gaps occur between neural arches
2-3 and the vertebral column, and between neu-
ral arches 3 and 4; these gaps are greatly reduced
in Thryssocypris (that between neural arches 3
and 4 is completely occluded). The element here
termed neural arch of centrum 2 is the anterior-
most supraneural according to Fink and Fink
(1981).

Fusion of centra 2 and 3 is possibly a primitive
character for all Cyprinidae. We propose a hy-
pothesis that this condition evolved only once
and that the mosaic distribution of cyprinid gen-

era with fused and with separate centra 2-3 is
due to repeated secondary failure to fuse and
reversion to the fused condition, rather than to
truly independent evolution of fusion between
these centra in phyletically diverse lines (the hy-
pothesis seemingly favored by Greenwood et al.
[1966:385] and Fink and Fink [1981:331]). Cen-
tra 2-3 are fused in the primitive siluroid Di-
plomystes, centra 2-4 and sometimes also 5 in
all other siluroids (Hassur 1 970). Whether fusion
of centra 2-3 evolved independently in siluroids
or indicates their phyletic relationship to cypri-
noids has not been resolved satisfactorily (com-
pare Roberts 1973 with Fink and Fink 1981).
The centra of the Weberian apparatus are all
separate in all characoids, gymnotoids, and gon-
orynchs. In any event this character is unlikely
to be relevant in considerations of the closer phy-
letic relationships of Thryssocypris.

PECTORAL GIRDLE (Fig. 1 6). — Pectoral girdle
morphologically generalized, slender, with a sin-
gle extrascapular (two extrascapulars in some
cyprinids), dorsal and anteroventral limbs of
cleithrum slender, postcleithrum moderately
elongate (slightly more elongate in T. tonlesa-
pensis than in T. smaragdinus). Coracoid with a
strongly developed lateral shelf, but not ventrally
expanded; coracoid foramen broad. (Coracoid is
ventrally expanded, sometimes enormously so,
in most chelines and cultrines. In some, coracoid
foramen reduced or even occluded.) In chelines
(Howes 1979), postcleithrum frequently (al-
ways?) arclike, very slender and elongate, and
with distal end projecting downwards, quite un-
like the postcleithrum in Thryssocypris. In
Thryssocypris the pectoral fin, although slightly
larger proportionately in T. tonlesapensis than
in T. smaragdinus, is not notably enlarged, nor
is the simple outer ray thickened or otherwise
modified. The pectoral axial flap (attached to the
postcleithrum) is simple, moderately elongate,
and not overlain by peculiarly shaped scales. In
all of these respects the pectoral fin of chelines
and cultrines tends to be modified.

PELVIC GIRDLE (Fig. 17). — Pubic bone deeply
bifurcate anteriorly (as in most cyprinoids); lat-
eral pubic projection nearly twice as long as me-
dial. Ischiac process highly modified, with ex-
tremely elongate anterior and posterior
projections. In T. smaragdinus anterior ischiac
projection about half as long as posterior pro-
jection; in T. tonlesapensis anterior projection

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 1

ischiac process

FIGURE 17. Thryssocypris smaragdinus, 53.2 mm, pelvic girdle (ventral view).

twice as long as posterior one, extending ante-
riorly as far as distal end of medial pubic pro-
jection. Origin of ischiac process opposite distal
end of pleural rib of vertebra number 9 in T.
smaragdinus, and of pleural rib of vertebra 1 1
or 1 2 in T. tonlesapensis.

AXIAL SKELETON. — Supraneurals 4-6 in num-
ber, commencing immediately posterior to neur-
al complex of Weberian apparatus (not greatly
enlarged or articulated to each other as in some
chelines). In T. smaragdinus vertebrae 5-20 and
in T. tonlesapensis 5-16 with enlarged prezyga-
pophyses. Thryssocypris smaragdinus with well-
developed dorsal ribs attached to anterior half
of centrum on vertebrae 1 1-24; T. tonlesapensis
without dorsal ribs (dorsal ribs unreported in any
Cyprinidae until now). Pleural ribs of vertebrae
5-10 with broad, laminar proximal ends at-
tached to comparably broad parapophyses; re-
maining pleural ribs with proximal ends slender,

weakly attached to slender parapophyses. In T.
smaragdinus proximal portion of pleural ribs on
vertebrae 6-7 with spurlike, posteriorly directed
projections (absent in T. tonlesapensis).

The counts of abdominal, caudal, and total
vertebrae in Thryssocypris (Table 1) are close or
identical to counts reported in several species of
Barilius by Howes (1980: table 1).

DORSAL AND ANAL FINS (Fig. 1 8). — Dorsal and
anal fin rays uniformly slender, nonserrate, first
two dorsal and first three anal fin rays simple.
Last complete pterygiophore bears two (counted
as one and a half) fully formed, branched rays;
bases of ray halves of anterior of these two rays
overlap and lie externally to ray halves of pos-
terior ray.

In the anal fin the bifurcate terminal radial (see
generic diagnosis) is the medial radial of the last
pterygiophore, which is complete. In the dorsal
fin the situation is more complicated, and the

fin ray

distal radial

proximal radial

FIGURE 18. Thryssocypris smaragdinus, 53.2 mm, posteriormost fin rays and pterygiophores in dorsal fin (lateral view above,
dorsal view below).

ROBERTS & KOTTELAT: THRYSSOCYPRIS, A NEW CYPRINID GENUS

157

homology of the bifurcate terminal radial with
the proximal, medial, and distal radials is un-
clear. In T. smaragdinus the bifurcate terminal
radial constitutes by itself the terminal pteryg-
iophore, articulating endochondrally only with
the medial radial of the preceding pterygiophore.
In T. tonlesapensis an additional element is pres-
ent ventral to the bifurcate terminal radial; the
dorsal margin of this element articulates with the
ventral margins of the bifurcate terminal radial
and of the preceding medial and proximal ra-
dials, and its homology also is unclear.

INTERMUSCULAR BONES. — Intermuscularbones
well-developed, epineurals extending entire length
of body, epipleurals commencing at vertebra
12. Proximal ends of anteriormost two or three
epineurals attached to exoccipital in a small
depression or fossa (shown in Fig. 8) near its
posterolateral margin and on a level with fora-
men magnum. Distal ends of posteriormost two
or three epineurals and epipleurals multifid
(mostly bifid or trifid). Epineurals and epipleur-
als forked anteriorly with expanded laminae pos-
teriorly.

In chelines and cultrines the anteriormost epi-
neurals tend to insert much higher on the cra-
nium (Howes 1979) than in Thryssocypris.

CAUDAL SKELETON (Fig. 19). — Generalized in
nearly all respects, with a single epural; one or
two uroneurals; parhypural fused to ural cen-
trum, with an exceptionally large, elongate hy-
purapophysis; six hypurals, separate from each
other, with hypurals 1-2 fused to ural complex
centrum.

In most lower teleosts with the primitive prin-
cipal caudal fin ray formula of 10 + 9, including
all characoids and all Cyprinidae, the caudal fin
skeleton tends to be highly conservative, retain-
ing a relatively generalized or primitive mor-
phology, including the primitive complement of
six hypurals (exclusive of the parhypural). There
is a maximum of three epurals, and perhaps in-
variably at least one epural. Presumably three is
the primitive number of epurals in teleosts in-
cluding ostariophysans, but reductions to two and
one and subsequent reversion to three seem to
have occurred repeatedly, particularly often in
ostariophysans including Cyprinidae. Thus the
single epural in Thryssocypris is not likely to be
helpful in assessing its relationships to other cyp-
rinids.

It should be noted that previous works on os-

uroneural

epural

urostyle
neural spine

ural centrum

hypurapophysis

parhypural

hemal spine

hypurals 1-6

FIGURE 19. Thryssocypris smaragdinus, 53.2 mm, caudal
fin skeleton (lateral view).

tariophysan caudal skeletons report seven hy-
purals (e.g., Weitzman 1962; Roberts 1969). Fol-
lowing Monod (1968) the element previously
considered as hypural one is here termed the
parhypural. The parhypural bears the hypura-
pophysis which serves as origin of the main
muscles for adduction of the upper caudal fin
lobe. Thus the hypurals formerly numbered 2-7
are now hypurals 1-6. This new nomenclature
has been used by most authors dealing with os-
tariophysan caudal skeletons subsequent to
Monod (1968) including Lundberg and Baskin
(1969) and Fink and Fink (1981).

CONCLUSION

The discovery of Thryssocypris provides
another example of the extraordinary diversity
of Cyprinidae, especially in Southeast Asia, and
of the close biogeographic relationship between
the Kapuas and Mekong basins.

Upon first examining Thryssocypris we thought
that it might be a specialized Barilius or at least
closely related to that genus. Lacking osteological
information on most other cyprinid genera we
are not prepared to discuss its relationships at
this time except to note that it does not seem to
be related to the chelines (sensu Howes 1979).
The specialized characters in Thryssocypris that
seem most unusual or highly derived and there-

158

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 1

fore likely to indicate phyletic relationships have
not been reported in any other cyprinids. These
include the peculiar morphology of ceratobran-
chials and absence of hypobranchials 2-3, shape
of jaw bones, elongate anterior and posterior pro-
jections of ischiac process, and bifid terminal
radials in median fins.

ACKNOWLEDGMENTS

For making available material of Thryssocy-
pris in their care we wish to thank J. Daget and
F. d'Aubenton and W. J. Rainboth. The study
was also facilitated in various ways by the fol-
lowing persons: G. J. Howes, Michael A. Hearne,
W. N. Eschmeyer, Daphne Dunn, and Vincent
Lee. The Kapuas ichthyological survey was
sponsored by the Smithsonian Tropical Research
Institute, Indonesian National Research Coun-
cil, and Museum Zoologicum Bogorense. This
paper is part of a project to report on the fishes
of the Kapuas basin supported by National Sci-
ence Foundation grant DEB77-24759.

LITERATURE CITED

FINK, S. V., AND W. L. FINK. 1981. Interrelationships of
ostariophysan fishes (Teleostei). Zool. J. Linn. Soc. 72(4):
297-353.

GOSLINE, W. A. 1975. The cyprinid dermosphenotic and the
subfamily Rasborinae. Occ. Pap. Mus. Zool. Univ. Michi-
gan, no. 673, 13 pp.

GREENWOOD, P. H., D. E. ROSEN, S. H. WEITZMAN, AND G. S.
MYERS. 1966. Phyletic studies of teleostean fishes, with a

provisional classification of living forms. Bull. Amer. Mus.

Nat. Hist. 131(4):339-456.
HASSUR, R. L. 1970. Studies on the osteology of catfishes,

order Siluriformes. Ph.D. dissertation, Stanford Univ., 133

PP-
HOWES, G. J. 1978. The anatomy and relationships of the

cyprinid fish Luciobrama macrocephalus (Lacepede). Bull.

Brit. Mus. (Nat. Hist.), zool. ser. 34(1): 1-64.
. 1979. Notes on the anatomy of Macrochirichthys

macrochirus (Valenciennes) 1844, with comments on the

Cultrinae (Pisces, Cyprinidae). Bull. Brit. Mus. (Nat. Hist.),

zool. ser. 36(3): 147-200.
. 1980. The anatomy, phylogeny and classification of

bariliine cyprinid fishes. Bull. Brit. Mus. (Nat. Hist.), zool.

ser. 37(3): 129-1 98.
LEKANDER, B. 1949. The sensory line system and the canal

bones in the head of some Ostariophysi. Acta Zool. 30:1-

131.

LUNDBERG, J. G., AND J. N. BASKIN. 1969. The caudal skel-
eton of the catfishes, order Siluriformes. Amer. Mus. Nov.

no. 2399, 49 pp.
MONOD,T. 1968. Le complex urophoredesteleosteens. Mem.

Inst. fr. Afr. noire 81:1-705.

RAMASWAMI, L. S. 1955. Skeleton of cyprinoid fishes in re-
lationship to phylogenetic studies: 7. The skull and Weberian

apparatus of Cyprininae (Cyprinidae). Acta Zool. 36:199-

242.
ROBERTS, T. R. 1969. Osteology and relationships of chara-

coid fishes, particularly the genera Hepsetus, Salminus, Ho-

plias, Ctenolucius, and Acestrorhynchus. Proc. California

Acad. Sci. 36(15):39 1-500.
. 1973. Interrelationships of ostariophysans, pp. 373-

395 in P. H. Greenwood, R. S. Miles, and C. Patterson (eds.),

Interrelationships of Fishes, London, Academic Press.
WEITZMAN, S. H. 1962. The osteology of Brycon meeki, a

generalized characid fish, with an osteological definition of

the family. Stanford Ichth. Bull. 8:1-77.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 12, pp. 159-177, 32 figs. July 12, 1984

STUDIES ON NEBRIINI (COLEOPTERA: CARABIDAE),

V. NEW NEARCTIC NEBRIA TAXA AND

CHANGES IN NOMENCLATURE

By
David 11. Kavanaugh

Department of Entomology, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118

ABSTRACT: This paper introduces new Nearctic Nebria taxa and changes in nomenclature. Names are pro-
vided for 10 new species and 3 new subspecies (type locality in parentheses): Nebria altisierrae (Olmsted
Point, Yosemite National Park, California), V. campbelli (Mount Baker, Whatcom County, Washington), V
wallowae (West Fork Wallowa River, Wallowa County, Oregon), N. Jeffrey! (South Fork McCoy Creek,
Harney County, Oregon), V. haida (Mount Needham, Graham Island, Queen Charlotte Islands, British
Columbia), N. louiseae (Skedans, Louise Island, Queen Charlotte Islands, British Columbia), N. gebleri
albimontis (Birch Creek, Mono County, California), N. labontei (West Fork Wallowa River, Wallowa County,
Oregon), V. calva (Mount Baldy, Apache County, Arizona), V. sierrablancae (Sierra Blanca, Lincoln County,
New Mexico), N. piute sevieri (Parowan Creek, Iron County, Utah), N. steensensis (South Fork McCoy
Creek, Harney County, Oregon), and N. trifaria pasquineli (Lefthand Creek, Boulder County, Colorado). For
each, diagnostic combination of characters and notes on geographical distribution are provided and distin-
guishing features are illustrated. Changes in status are proposed for the following names (second name in
each pair considered valid): Nebria intermedia Van Dyke = N. crassicornis intermedia Van Dyke; N. sonorae
Kavanaugh = N. acuta sonorae Kavanaugh; N.fragilis Casey = N. arkansana fragilis Casey; N. trifaria piute
Erwin and Ball = N. piute piute Erwin and Ball; and N. trifaria utahensis Kavanaugh = N. piute utahensis
Kavanaugh. New synonymies proposed include: Nebria arkansana uinta Kavanaugh = N. arkansana fragilis
Casey; N. fragilis teewinot Kavanaugh = TV. arkansana fragilis Casey; and N. trifaria tetonensis Erwin and
Ball = N. trifaria trifaria LeConte.

INTRODUCTION imens and data have been acquired. Study of this
For several years, I have been working on a new material has revealed 1 3 hitherto unknown
monographic treatment of genus Nebria Latreille taxa ( 1 0 species and 3 subspecies) as well as sev-
for the Nearctic Region. During that time, I have eral nomenclatural problems with previously de-
provided names for several new species and nu- scribed taxa.

merous new subspecies, designated lectotypes, The purpose of this report, which serves as a
and proposed certain nomenclatural changes final presentation of nomenclatural matters pre-
(Ka vanaugh 1979 and 1981). Validation of these liminary to submission of the monographic treat-
new names and clarification of the status of ex- ment, is to provide names for the new taxa and
isting names were needed to permit their proper to present formally the needed nomenclatural
use in various other reports by the author and changes. The latter include both new synonymies
several colleagues. Since 1981, additional spec- and other changes in status of names. As before,

[159]

160

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12

these names are needed immediately for use in
other manuscripts; and data and discussions pre-
sented for each name are limited to little more
than the minimum required by the International
Code of Zoological Nomenclature. More detailed
information for all taxa, including those pre-
sented here as new, will be provided in the mono-
graph.

MATERIALS AND METHODS

This report is based on examination of 1 136
adult Nebria specimens, representing taxa de-
scribed here as new, and over 80,000 additional
specimens, representing previously described
Nebria taxa, used for comparative purposes.
Recognition of new synonymies and changes in
status of taxa are based on study of this total
specimen resource pool.

Following is a list of acronyms used in the text.
These refer to collections from which specimens
have been received and/or in which paratype
specimens have been deposited. Curators re-
sponsible for collections during the course of my
study are also listed; and I here acknowledge with
sincere thanks their assistance in providing spec-
imens for study on loan.

ANSP— Academy of Natural Sciences, Philadelphia, Pennsyl-
vania 19103; W. W. Moss.

BCPM— British Columbia Provincial Museum, Victoria, Brit-
ish Columbia V8V 1X4; R. A. Cannings.

CArm-C. Armin, 191 West Palm Avenue, Reedley, Califor-
nia 93654 (specimens deposited in CAS).

CAS— California Academy of Sciences, San Francisco, Cali-
fornia 941 18; D. H. Kavanaugh, H. B. Leech.

CNC— Canadian National Collection of Insects, Biosystem-
atics Research Institute, Ottawa, Ontario K1A OC6; R. de
Ruette, A. Smetana.

CUB— University of Colorado, Boulder, Colorado 80302; H.
Rodeck.

CUIC-Cornell University, Ithaca, New York 14850; L. L.
Pechuman.

DEUN— University of Nebraska, Lincoln, Nebraska 68503;
B. C. Ratcliffe.

EAMa— E. A. Martinko, University of Kansas, Lawrence,
Kansas 66044 (specimens deposited in CAS).

FMNH— Field Museum of Natural History, Chicago, Illinois
60605; H. Dybas, L. Watrous.

JRLa-J. R. La Bonte, 7 10 NW 1 1th, Corvallis, Oregon 97330.

KSUC— Kansas State University, Manhattan, Kansas 66502;
H. D. Blocker.

LACM— Los Angeles County Museum of Natural History, Los
Angeles, California 90007; C. L. Hogue.

MCZ— Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge, Massachusetts 02138; J. F. Lawrence, A.
F. Newton, Jr.

MSU— Michigan State University, East Lansing, Michigan
48823; R. L. Fischer.

OSUC-Ohio State University, Columbus, Ohio 43210; C. A.
Triplehorn.

PJJo-P. J. Johnson, 1408 28th Street SE, Auburn, Washing-
ton 98002.

QCIM— Queen Charlotte Islands Museum, Skidegate, British
Columbia VOT ISO; N. Gessler and T. Gessler.

ROM— Royal Ontario Museum, Toronto, Ontario M5S 2C6;
G. B. Wiggins.

RTBe-R. T. Bell, University of Vermont, Burlington, Ver-
mont 05401.

UAFA— University of Arkansas, Fayetteville, Arkansas 72701;
R.T.Allen.

U ASM— University of Alberta, Strickland Museum, Edmon-
ton, Alberta T6G 2E3; G. E. Ball.

UMMZ— University of Michigan, Ann Arbor, Michigan 48104;
R. D. Alexander.

USNM— United States National Museum, Smithsonian Insti-
tution, Washington, D.C. 20560; T. L. Erwin.

UWEM— University of Wisconsin, Madison, Wisconsin 53706;
L. J. Bayer.

ZMLS— Zoological Institute, University of Lund, Lund, Swe-
den; C. H. Lindroth, R. Danielsson.

Methods used in the present study, including
measurement and dissection techniques and cri-
teria for ranking taxa as species or subspecies,
have been described in a previous paper (Ka-
vanaugh 1979).

NEW NEBRIA SPECIES AND SUBSPECIES

The order of presentation of new species and
subspecies below reflects a new classification of
Nearctic Nebria, which will be provided in a
monographic treatment of the genus now in
preparation for publication. Except as noted, all
specimens listed (by locality) in sections about
geographical distribution have been designated
as paratypes.

Nebria altisierrae, new species

(Figures 2, 15,31)

Nebria virescens; KAVANAUGH 1978:345 (in part).

HOLOTYPE, (J, in CAS, labelled: "U.S.A., California, Yosem-
ite N. P., Sierra Nevada, Olmsted Point, 1.5 mi. sw. Tenaya
Lk. on Tioga Rd., 2560m, 8 Nov. 1976 D. H. Kavanaugh"/
"D. H. Kavanaugh Collection" [orange label]/"Holotype Ne-
bria altisierrae n. sp. del. D. H. Kavanaugh 1983" [red label]/
"California Academy of Sciences Type No. 14338." PARATYPES:
five (four <J and one 9), also deposited in CAS.

TYPE-LOCALITY.— Olmsted Point, 2560 m, Sierra Nevada,
Yosemite National Park, California.

DIAGNOSTIC COMBINATION.— Head uniformly
dark, without pale spots on vertex; labium with
paraglossae distinct as short, pointed lateral lobes
on apical margin of ligula; pronotum (Fig. 2)
semiovoid, relatively short and moderately wide,
ratio pronotal width to elytral width less than or
equal to 0.75, basal sinuation of lateral margin

KAVANAUGH: NEW NEARCTIC NEBRIA

161

absent or short and very shallow, basal angles
not or only moderately denticulate, midlateral
seta present; elytra with silhouette subrectan-
gular, not narrowed basally, humeri (Fig. 1 5) not
or only faintly carinate; metepisternum impunc-
tate; hind tarsus with all tarsomeres glabrous
dorsally, fourth tarsomere truncate ventrally with
medial and lateral apicoventral setae symmet-
rical in length and position; specimen from lo-
cality in Sierra Nevada of California (Fig. 31).

DERIVATION OF TAXON NAME.— The species
epithet is a combination of the Latin word for
"high" (—altus) and a shortened form of "Sierra
Nevada," in reference to the High Sierra region
inhabited by members of this species.

GEOGRAPHICAL DISTRIBUTION. — Figure 31.
Known only from high elevations in the Sierra
Nevada of California, from Tuolumne County
south to Sequoia National Park. I have studied
specimens from the following localities:

United States of America

CALIFORNIA: Sequoia National Park, Alta Meadow ([2740
m]) [Aug.] (2; CAS); Tuolumne County, Blue Canyon Lake
([3110 m]) [July] (1; CAS); Yosemite National Park, Olmsted
Point (1.5 miles SW of Tenaya Lake [2560 m]) [Nov.] (3; CAS).

Nebria campbelli, new species

(Figures 3, 16, 31)

Nebria virescens\ KAVANAUGH 1978:345 (in part).

HOLOTYPE, 6, in CAS, labelled: "U.S.A., Wash., Whatcom
Co., Cascade Range, ne. slope Mt. Baker, Kulshan Ridge,
1460m-1520m, 11 Aug. 74 D. H. Kavanaugh'V "D. H. Ka-
vanaugh Collection" [orange label]/ "Holotype Nebria camp-
belli n. sp. del. D. H. Kavanaugh 1983" [red label]/ "California
Academy of Sciences Type No. 14339." PARATYPES: two (one
i and one 9) deposited in CAS and CNC.

TYPE-LOCALITY. — Mount Baker, 1460-1520 m, Cascade
Range, Whatcom County, Washington.

DIAGNOSTIC COMBINATION. — Head uniformly
dark, without pale spots on vertex; labium with
paraglossae distinct as short, pointed lateral lobes
on apical margin of ligula; pronotum (Fig. 3)
semiovoid, relatively long and narrow, basal sin-
uation of lateral margin short and moderately
deep, basal angles markedly denticulate laterally,
midlateral seta present; elytra with silhouette
narrowed basally, nearly subovoid, humeri (Fig.
16) markedly carinate; metepisternum impunc-
tate; hind tarsus with all tarsomeres glabrous
dorsally, fourth tarsomere truncate ventrally with
medial and lateral apicoventral setae symmet-
rical in length and position.

DERIVATION OF TAXON NAME. — I take plea-

sure in naming this species in honor of my friend
and colleague J. Milton Campbell, who collected
the only known female specimen of this species.
GEOGRAPHICAL DISTRIBUTION. — Figure 31.
Known only from the northern end of the Cas-
cade Range and its flanking ranges, from Man-
ning Provincial Park in southern British Colum-
bia to Mount Baker in northern Washington. I
have studied specimens from the following lo-
calities:

Canada

BRITISH COLUMBIA: Manning Provincial Park (Three
Brothers Mountain [2130 m]) [July] (1; CNC).

United States of America

WASHINGTON: Okanogan County, Pasayten Wilderness
(Bunker Hill Lookout [2120 m]) (1; CAS); Whatcom County,
Mount Baker (NE slope on Kulshan Ridge [1520 m]) [Aug.]
(1;CAS).

Nebria wallowae, new species

(Figures 4, 17, 24, 31)

HOLOTYPE, <5, in CAS, labelled: "U.S.A., Oregon, Wallowa
County, Wallowa Mts., West Fork Wallowa River, 2070-
2130m, 10 July 1982 Stop #82-16 D.H. & J. L. Kavanaugh
colls."/ "D. H. Kavanaugh Collection" [orange label]/ "Ho-
lotype Nebria wallowae n. sp. del. D. H. Kavanaugh 1983"
[red label]/ "California Academy of Sciences Type No. 1 4347."
PARATYPES: 87 (36 $ and 5 1 S), deposited in CAS, CNC, JRLa,
UASM, and USNM.

TYPE-LOCALITY. - West Fork Wallowa River, 2070-2 1 30 m,
Wallowa Mountains, Wallowa County, Oregon.

DIAGNOSTIC COMBINATION. — Pronotum (Fig.
4) with lateral explanation broad throughout,
basal sinuation of lateral margin moderately deep,
basal angles rectangular or slightly acute, mod-
erately projected posteriorly, midlateral seta ab-
sent; elytra with silhouette subovoid, slightly
narrowed basally, humeral angles not markedly
rounded or obtuse, humeral carinae moderately
developed, slightly projected anteriorly; hind-
wing full-sized, with reflexed apex distal to stig-
ma; hind coxae bi- or plurisetose basally; middle
tibiae dorsally concave or sulcate (at least near
middle), with brush of dorsal setae moderately
and densely developed; third to fifth visible ab-
dominal sterna each with two or more pairs of
posterior paramedial setae; median lobe of male
(Fig. 1 7) long, very slender; bursa copulatrix of
female (Fig. 24) with very small bursal sclerite;
specimen from locality in Wallowa Mountains
of Oregon (Fig. 31).

DERIVATION OF TAXON NAME.— This species
is named for the Wallowa Mountains.

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12

GEOGRAPHICAL DISTRIBUTION. — Figure 31.
Known only from the Wallowa Mountains of
northeastern Oregon. I have studied specimens
from the following localities:

United States of America

OREGON: Wallowa County, Glacier Lake (meadow below
[2440 m]) [Sep.] (2; JRLa), Mirror Lake area ([2350 m]) [July]
(5; CNC), West Fork Wallowa River (0.25 to 0.5 miles NE of
Frazier Lake [2040-2190 m], and small side stream [1980 m])
[July] (81; CAS, JRLa).

Nebria jeffreyi, new species

(Figures 5, 31)

HOLOTYPE, 3, in CAS, labelled: "U.S.A., Oregon, Harney
County, Steens Mountains, South Fork McCoy Creek, 2290-
2560m, 14 July 1982, Stop #82-22, D. H. & J. L. Kavanaugh
colls."/ "D. H. Kavanaugh Collection" [orange label]/ "adult
specimen used in laboratory rearing of immature stages"/ "Ho-
lotype Nebria jeffreyi n. sp. det. D. H. Kavanaugh 1983" [red
label]/ "California Academy of Sciences Type No. 14342."
PARATYPES: 35 (1 1 $ and 24 9), deposited in CAS, JRLa, PJJo,
UASM, and USNM.

TYPE-LOCALITY. -South Fork McCoy Creek, 2390-2560 m,
Steens Mountains, Harney County, Oregon.

DIAGNOSTIC COMBINATION.— Pronotum (Fig.
5) with lateral explanation broad throughout,
apical angles very broad and rounded, basal an-
gles rectangular, basal sinuation of lateral margin
moderately deep, midlateral seta absent; elytra
with silhouette subrectangular, not narrowed ba-
sally, humeral angles not markedly rounded or
obtuse, humeral carinae absent or only slightly
developed, not projected anteriorly; hind coxae
bi- or plurisetose basally; middle tibiae dorsally
concave or sulcate (at least near middle), with
brush of dorsal setae moderately and densely de-
veloped; third to fifth visible abdominal sterna
each with two or more pairs of posterior para-
medial setae; specimen from locality in Steens
Mountains of Oregon (Fig. 31).

DERIVATION OF TAXON NAME.— I take great
pleasure in naming this species in honor of my
son, Jeffrey L. Kavanaugh, who assisted me in
collecting the first known specimens of this
species.

GEOGRAPHICAL DISTRIBUTION. — Figure 31.
Known only from the Steens Mountains of
southcentral Oregon. I have studied specimens
from the following localities:

United States of America

OREGON: Harney County, Little Blitzen River ([2560 m])
[July] (10; PJJo), South Fork McCoy Creek ([2390-2560 m])
[July] (26; CAS, JRLa).

Nebria haida, new species

(Figures 6, 31)

HOLOTYPE, 3, in CAS, labelled: "Canada, British Columbia,
Queen Charlotte Islands, Graham Island, 1.8 km N of Mt.
Needham, 700m-780m, 18 July 1981, Stop #81-37, D. H.
Kavanaugh collector"/ "D. H. Kavanaugh Collection" [orange
label]/ "Queen Charlotte Islands Expedition— 1981" [row of
asterisks] "D. H. Kavanaugh Calif. Acad. Sciences"/ "Holo-
type Nebria haida n. sp. det. D. H. Kavanaugh 1983" [red
label]/ "California Academy of Sciences Type No. 14341."
PARATYPES: 151 (68 $ and 83 2), deposited in BCPM, CAS,
CNC, QCIM, UASM, and USNM).

TYPE-LOCALITY.— 1.8 km N of Mount Needham, 700-780
m, Graham Island, Queen Charlotte Islands, British Columbia.

DIAGNOSTIC COMBINATION. — Size medium:
standardized body length of male 9.0 to 10.5
mm, of female 9.4 to 10.8 mm; head relatively
large and wide; pronotum (Fig. 6) relatively long
and slender, with lateral explanation broad
throughout, apical angles short, relatively broad
and slightly rounded, basal angles rectangular,
basal sinuation of lateral margin long, moder-
ately deep, midlateral seta absent; elytra with
silhouette subovoid, long and slender, distinctly
narrowed basally, humeral angles not markedly
rounded or obtuse, humeral carinae absent or
only slightly developed, not projected anteriorly,
intervals markedly convex; legs long, slender,
femora and tibiae piceous; hind coxae bi- or
plurisetose basally; middle tibiae dorsally con-
cave or sulcate (at least at middle), with brush
of dorsal setae moderately and densely devel-
oped; third to fifth visible abdominal sterna each
with two or more pairs of posterior paramedial
setae; specimen from locality in Queen Charlotte
Islands, British Columbia (Fig. 31), from above
treeline in alpine area.

DERIVATION OF TAXON NAME.— This species
is named in honor of the Haida people, tradi-
tional inhabitants of the Queen Charlotte Ar-
chipelago.

GEOGRAPHICAL DISTRIBUTION. — Figure 31.
Known only from high elevations in the Queen
Charlotte Islands, British Columbia. I have stud-
ied specimens from the following localities:

Canada

BRITISH COLUMBIA: Queen Charlotte Islands: Graham Is-
land, Mount Needham ( 1 .0 km [790-9 1 0 m] and 1 .8 km [700-
780 m] N) [July] (148; CAS); Moresby Island, Mount Moresby
(northwest-facing cirque [910-1070 m]) [July] (4; CAS).

Nebria louiseae, new species

(Figures 7, 31)

HOLOTYPE, 6, in CAS, labelled: "B. C., Q. C. I. Louise Is.,

KAVANAUGH: NEW NEARCTIC NEBRIA

163

Skedans 1 1. VI. 1981 R. A. Cannings"/ "Holotype Nebria lou-
iseae n. sp. del. D. H. Kavanaugh 1983" [red label]/ "California
Academy of Sciences Type No. 15005." PARATYPES: four 9,
deposited in BCPM, CAS, and CNC.

TYPE- LOCALITY.— Skedans, Louise Island, Queen Charlotte
Islands, British Columbia.

DIAGNOSTIC COMBINATION.— Size large: stan-
dardized body length of male 10.4 mm, of female
10.6 to 1 1.0 mm; head relatively large and wide;
pronotum (Fig. 7) relatively long and slender,
with lateral explanation broad throughout, apical
angles moderate in length, relatively narrow and
pointed, basal angles rectangular, basal sinuation
of lateral margin long, moderately deep, midlat-
eral seta absent; elytra with silhouette subovoid,
long and slender, distinctly narrowed basally, hu-
meral angles not markedly rounded or obtuse,
humeral carinae absent or only slightly devel-
oped, not projected anteriorly, intervals mark-
edly convex; legs long, slender, femora and tibiae
piceous; hind coxae bi- or plurisetose basally;
middle tibiae dorsally concave or sulcate (at least
at middle), with brush of dorsal setae moderately
and densely developed; third to fifth visible ab-
dominal sterna each with two or more pairs of
posterior paramedial setae; specimen from lo-
cality in Queen Charlotte Islands, British Colum-
bia (Fig. 31), from upper sea beach area.

DERIVATION OF TAXON NAME.— This species
is named for Louise Island, on which the type
locality is found.

GEOGRAPHICAL DISTRIBUTION.— Figure 31. At
present known only from Louise Island, Queen
Charlotte Islands. I have studied specimens from
the following locality:

Canada

BRITISH COLUMBIA: Queen Charlotte Islands: Louise Island,
Skedans [June] (5; BCPM).

Nebria gebleri albimontis, new subspecies

(Figures 1, 8, 18, 25, 32)

HOLOTYPE, 3, in CAS, labelled: "U.S.A., California, Mono
County, White Mts., Birch Creek, 3290m-3410m, 8 July 1980
D. Giuliani collector"/ "Collection of California Academy of
Sciences, San Francisco, Calif."/ "Holotype Nebria gebleri al-
bimontis n. ssp. det. D. H. Kavanaugh 1983" [red label]/ "Cal-
ifornia Academy of Sciences Type No. 14340." PARATYPES:
four (two <5 and two 9), also deposited in CAS.

TYPE-LOCALITY. — Birch Creek, 3290-3410 m, White Moun-
tains, Mono County, California.

DIAGNOSTIC COMBINATION.— Head dark, with
a pair of pale paramedial spots on vertex; anten-
nal scape (Fig. 1) short, moderately thick; prono-

tum (Fig. 8) with basal angles markedly acute,
distinctly divergent posteriorly, margination of
apical angles and anterior one-third of lateral
margin very narrow, midlateral and basolateral
setae present; elytra without metallic reflection,
elytral silhouette subrectangular, hindwing full-
sized; median lobe of male (Fig. 1 8) with pre-
apical area markedly bulbous left dorsolaterally;
bursa copulatrix of female (Fig. 25) with sper-
mathecal chamber small, narrow in dorsal as-
pect; specimen from locality in White Mountains
of California (Fig. 32).

DERIVATION OF TAXON NAME.— The subspe-
cific epithet is a combination of the Latin words
for "white" (=albus) and "mountain" (=mons),
in reference to the White Mountains.

GEOGRAPHICAL DISTRIBUTION. — Figure 32.
Known only from the type locality in the White
Mountains of eastern California. I have studied
specimens from the following locality:

United States of America

CALIFORNIA: Mono County, Birch Creek ([3290-3410 m])
[July] (5; CAS).

Nebria labontei, new species

(Figures 9, 32)

HOLOTYPE, S, in CAS, labelled: "U.S.A., Oregon, Wallowa
County, Wallowa Mts., West Fork Wallowa River, 2040-
2190m, 11 July 1982, D. H. & J. L. Kavanaugh Stop #82-
17"/ "D. H. Kavanaugh Collection" [orange label]/ "adult
specimen used in laboratory rearing of immature stages"/ "Ho-
lotype Nebria labontei n. sp. det. D. H. Kavanaugh 1983" [red
label]/ "California Academy of Sciences Type No. 14343."
PARATYPES: 57 (29 6 and 28 9), deposited in CAS, JRLa, UASM,
and USNM.

TYPE-LOCALITY. - West Fork Wallowa River, 2040-2 190m,
Wallowa Mountains, Wallowa County, Oregon.

DIAGNOSTIC COMBINATION.— Size very large:
standardized body length of male greater than
1 1.5 mm, of female greater than or equal to 12.0
mm; head moderate in size, dark, with a pair of
pale paramedial spots on vertex; pronotum (Fig.
9) broad, markedly cordate, midlateral and ba-
solateral setae present; elytra with brilliant red
metallic reflection, elytral silhouette distinctly
subovoid, narrowed basally with lateral margins
distinctly rounded, intervals flat; specimen from
locality in Wallowa Mountains of Oregon (Fig.
32).

DERIVATION OF TAXON NAME.— I am pleased
to name this species in honor of my friend and
fellow collector, James R. LaBonte, who col-
lected the first known specimen of this extraor-
dinary species.

164

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12

GEOGRAPHICAL DISTRIBUTION. — Figure 32.
Known only from the Wallowa Mountains of
northeastern Oregon. I have studied specimens
from the following localities:

United States of America

OREGON: Wallowa County, Glacier Lake (meadow below
[2440 m]) [Sep.] (1; JRLa), West Fork Wallowa River (0.25 to
0.5 miles NE of Frazier Lake [2040-2190 m]) [July] (57; CAS,
JRLa).

Nebria calva, new species

(Figures 10, 19, 26, 32)

HOLOTYPE, 3, in CNC, labelled: "Ariz: Apache Co Mt. Baldy,
10-1 1000' SW of Springerville 13.VII.79, S & J Peck spruce-
fir forest"/ "Holotype Nebria calva n. sp. det. D. H. Kavanaugh
1983" [red label]. PARATYPES: 10 (7 3 and 3 2), deposited in
CAS and CNC.

TYPE-LOCALITY.— Mount Baldy (SW of Springerville), 3050-
3350 m, Apache County, Arizona.

DIAGNOSTIC COMBINATION.— Head moderate
in width and size, dark, with a pair of pale par-
amedial spots on vertex; pronotum (Fig. 10) with
midlateral and basolateral setae present; elytra
with faint but distinct metallic (violet) reflection,
elytral silhouette subovoid, markedly narrowed
basally, intervals moderately flat; median lobe
of male (Fig. 1 9) with apex straight in ventral
aspect; bursa copulatrix of female (Fig. 26) with
very small bursal sclerite; specimen from locality
in eastcentral Arizona (Fig. 32).

DERIVATION OF TAXON NAME.— The species
epithet is formed from the Latin word for "bald"
(=calvus), in reference to the type locality, Mount
Baldy.

GEOGRAPHICAL DISTRIBUTION. — Figure 32.
Known only from Mount Baldy in eastern Ari-
zona. I have studied specimens from the follow-
ing locality:

United States of America

ARIZONA: Apache County, Mount Baldy (SW of Springerville
[3050-3350 m]) [July] (11; CNC).

Nebria sierrablancae, new species

(Figures 11,20,27, 32)

Nebria trifaria catenata; KAVANAUGH 1978:431 (in part).

HOLOTYPE, 3, in CNC, labelled: "N. M. Lincoln Co. Sierra
Blanca 10500' 18. VII. 1969 A. Smetana"/ "Holotype Nebria
sierrablancae n. sp. det. D. H. Kavanaugh 1983" [red label].
PARATYPES: 25 (10 3 and 152), deposited in CAS, CNC, and
CUIC.

TYPE-LOCALITY.— Sierra Blanca, 3200 m, Lincoln County,
New Mexico.

DIAGNOSTIC COMBINATION. — Body color uni-
formly rufous; head relatively large in relation

to pronotum, with a pair of pale paramedial spots
on vertex; pronotum (Fig. 1 1) with lateral margin
markedly sinuate basally, slightly to moderately
angulate at middle, apical angles narrow and
bluntly pointed, midlateral and basolateral setae
present; elytra with faint metallic (violet) reflec-
tion, elytral silhouette subovoid, narrowed ba-
sally, intervals moderately flat; median lobe of
male (Fig. 20) with apex deflected left laterally
in ventral aspect; bursa copulatrix of female as
in Fig. 27; specimen from locality in Capitan
Mountains or Sierra Blanca of central New Mex-
ico (Fig. 32).

DERIVATION OF TAXON NAME.— This species
is named for Sierra Blanca, the type locality.

GEOGRAPHICAL DISTRIBUTION. — Figure 32.
Known only from Sierra Blanca and the Capitan
Mountains of central New Mexico. I have stud-
ied specimens from the following localities:

United States of America

NEW MEXICO: Lincoln County, Capitan (1; CUIC), Sierra
Blanca (Sierra Blanca Ski Area [3200-3510 m]) [July] (25;
CNC).

Nebria piute sevieri, new subspecies

(Figures 12,21,28, 32)

Nebria trifaria trifaria, auctorum— ERWIN AND BALL 1972:93
(in part)— KAVANAUGH 1978:430 (in part).

HOLOTYPE, 3, in CAS, labelled: "U.S., Utah, Iron Co., Mar-
kagunt Plateau, 13.5 mi. s. Parowan, Hwy. 143, Parowan Cr.,
9200', 21 June 71 D H Kavanaugh & E A Martinko"/ "D. H.
Kavanaugh Collection" [orange label]/ "71-224" [orange la-
bel]/ "Holotype Nebria piute sevieri n. ssp. det. D. H. Kava-
naugh 1983" [red label]/ "California Academy of Sciences Type
No. 14344." PARATYPES: 230 (107 3 and 123 2), deposited in
ANSP, CAS, CNC, KSUC, MCZ, OSUO, UASM, and USNM.
All specimens studied have been designated as paratypes ex-
cept for the single (female) specimen from Clay Springs, Navajo
County, Arizona. At present, I believe that this specimen is
mislabelled. However, form of the bursa copulatrix in this
specimen diners markedly from that in other females of N.
piute sevieri. This may be a teratological example, or, if the
specimen is correctly labelled, it may represent a distinct form
not yet adequately sampled. My identification of the specimen
as belonging to this subspecies is therefore tentative, pending
additional fieldwork in Arizona.

TYPE- LOCALITY. — Parowan Creek (1 3.5 miles S of Parowan),
2800 m, Markagunt Plateau, Iron County, Utah.

DIAGNOSTIC COMBINATION.— Body color uni-
formly rufopiceous or black; head moderate in
width and size in relation to pronotum, with a
pair of pale paramedial spots on vertex; antennal
scape markedly narrowed basally; pronotum (Fig.
12) with lateral margin moderately sinuate ba-
sally, rounded at middle, apical angles relatively

KAVANAUGH: NEW NEARCTIC NEBRIA

165

broad and rounded, midlateral and basolateral
setae present; elytra without metallic reflection,
elytral silhouette elongate, subovoid or nearly
ovoid, narrowed basally, intervals moderately
flat; median lobe of male (Fig. 21) very thick
basal to apical orifice, with apex moderate in
length and broad in lateral aspect, deflected left
laterally in ventral aspect; bursa copulatrix of
female (Fig. 28) with bursal sclerite large, narrow
in dorsal aspect; specimen from locality in south-
western Utah, east of Tushar Mountains and
Midget Crest and west of Henry Mountains (Fig.
32).

DERIVATION OF TAXON NAME.— This subspe-
cies is named for the Sevier River and Sevier
Plateau, important physiographic features of the
region occupied by members of this species.

GEOGRAPHICAL DISTRIBUTION. — Figure 32.
Known at present only from the montane region
between Salina, Utah, and Cedar Breaks Na-
tional Monument and east to the Boulder Moun-
tains. I have studied specimens from the follow-
ing localities:

United States of America

UTAH: Garfield County, Cottonwood Peak (38.5 miles SW
of Antimony at Cottonwood Creek [2440 m]) [July] (72; CAS),
Mount Dutton (23.4 miles SW of Antimony at North Fork
Deep Creek [3120 m]) [July] (18; CAS); Iron County, Cedar
Breaks National Monument ([3200 m]) [June-Aug.] (36; CAS,
OSUC), Cedar Canyon (Coal Creek [2650 m]) [June] (6; CAS),
The Mammoth ([3050 m]) [July] (17; ANSP, CAS, CNC, KSUC,
MCZ, UASM, USNM), Parowan Creek (13.5 miles S of Par-
owan [2800 m]) [June] (10; CAS); Kane County, Long Valley
Junction [Aug.] (12; CAS); Sevier County, Monroe Peak (8.6
[2640 m] and 12.4 [2990 m] miles SE of Monroe) [July] (3;
CAS), Mount Marvine (0. 1 miles N of Johnson Valley Res-
ervoir at Sevenmile Creek [2590 m]) [Aug.] (10; CAS); Wayne
County, Bluebell Knoll (31 miles S of Torrey [2440-3050 m])
[July] (46; CAS, USNM).

Doubtful Records:

UNITED STATES OF AMERICA— ARIZONA: Navajo County, Clay
Springs [Sep.] (1; CAS).

Nebria steensensis, new species

(Figures 12,22,29, 32)

HOLOTYPE, S, in CAS, labelled: "U.S.A., Oregon, Harney
County, Steens Mountains, South Fork McCoy Creek, 2390-
2560m, 14 July 1982, Stop #82-22 D. H. & J. L. Kavanaugh
colls."/ "D. H. Kavanaugh Collection" [orange label]/ "Ho-
lotype Nebria steensensis n. sp. det. D. H. Kavanaugh 1983"
[red label]/ "California Academy of Sciences Type No. 14345."
PARATYPES: 106 (52 6 and 54 2), deposited in CAS, JRLa, PJJo,
UASM, and USNM.

TYPE-LOCALITY. -South Fork McCoy Creek, 2390-2560 m,
Steens Mountains, Harney County, Oregon.

DIAGNOSTIC COMBINATION. — Body color uni-
formly black; head moderate in width and size
in relation to pronotum, with a pair of pale par-
amedial spots on vertex; antennal scape slightly
narrowed basally; pronotum (Fig. 1 3) with lateral
margin very deeply sinuate basally, rounded at
middle, apical angles relatively broad and round-
ed, basal angles rectangular, midlateral and ba-
solateral setae present; elytra without metallic
reflection, elytral silhouette subovoid, narrowed
basally, intervals moderately flat; median lobe
of male (Fig. 22) moderate in thickness basal to
apical orifice, with distinct, projected ridge on
right lateral surface, apex deflected left laterally
in ventral aspect; bursa copulatrix of female (Fig.
29) with bursal sclerite small; specimen from lo-
cality in Steens Mountains of Oregon (Fig. 32).

DERIVATION OF TAXON NAME.— The species is
named for the Steens Mountains.

GEOGRAPHICAL DISTRIBUTION. — Figure 32.
Known only from the Steens Mountains of south-
central Oregon. I have studied specimens from
the following localities:

United States of America

OREGON: Harney County, Fish Creek Gorge ([2 1 90 m]) [July]
(1; PJJo), Kiger Headwall ([2680 m]) [Aug.] (1; PJJo), Little
Blitzen River ([2560 m]) [July] (16; PJJo), Pate Lake ([2260
m]) [July] (1; PJJo), South Fork McCoy Creek ([2390-2560
m]) [July] (88; CAS, JRLa).

Nebria trifaria pasquineli, new subspecies

(Figures 14, 23, 30, 32)

Nebria trifaria coloradensis; ERWIN AND BALL 1972:96 (in part).
Nebria trifaria trifaria; KAVANAUGH 1978:430 (in part).

HOLOTYPE, S, in CAS, labelled: "Lefthand Cr., 5 mi. E. Ward,
Colo. Bould. Co. 20 July 68"/ "D. H. Kavanaugh Collection"
[orange label]/ "Holotype Nebria trifaria pasquineli n. ssp. det.
D. H. Kavanaugh 1983" [red label]/ "California Academy of
Sciences Type No. 14346." PARATYPES: 407 (224 3 and 183
9), deposited in ANSP, CArm, CAS, CNC, CUB, DEUN, EAMa,
FMNH, KSUC, LACM, MCZ, MSU, ROM, RTBe, UAFA,
UASM, UMMZ, USNM, UWEM, and ZMLS.

TYPE-LOCALITY.— Lefthand Creek (5 miles E of Ward), Front
Range, Boulder County, Colorado.

166

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12

lateral setae present; elytra without metallic re-
flection, moderately dull, microsculpture
moderately impressed, elytral silhouette sub-
ovoid, elongate, narrowed basally, intervals
moderately flat, fifth interval interrupted, mod-
erately or markedly catenate (with 3 to 10 ca-
tenations, restricted to apical one half or also on
basal one-half of interval); median lobe of male
(Fig. 23) slender basal to apical orifice, without
ridge on right lateral surface, apex moderate in
thickness and length and bent dorsally in lateral
aspect, deflected left laterally in ventral aspect,
apical orifice short, slightly constricted; bursa
copulatrix of female (Fig. 30) with bursal sclerite
small, narrow; specimen from locality in eastern
ranges of Southern Rocky Mountains of central
or northcentral Colorado or southeastern Wyo-
ming (Fig. 32).

DERIVATION OF TAXON NAME.— This subspe-
cies is named for Pasquinel, the enigmatic, in-
domitable French trapper in James A. Miche-
ner's epic novel, Centennial. The mountains
traveled and loved by this fictional character are
within the geographical range of this taxon and,
in fact, include the type locality.

GEOGRAPHICAL DISTRIBUTION. — Figure 32.
Known only from the easternmost ranges of the
Southern Rocky Mountains, from the Medicine
Bow and Sierra Madre Ranges of southeastern
Wyoming south to the Rampart Range of south-
central Colorado. I have studied specimens from
the following localities:

United States of America

COLORADO: (3; ANSP, KSUC, LACM); Boulder County,
[Aug.] (1; CUB), Arapahoe Pass ([2740-3050 m]) [Aug.] (5;
RTBe), Jenny Lake ([3200 m]) [July] (1; CArm), Lake Isabelle
([3170 m]) [July] (7; CArm), Jasper Lake ([3260 m]) [Aug.] (1;
CArm), Lefthand Creek (5 miles E of Ward [2530 m]) [July-
Aug.] (80; CArm, CAS), Little Royal Gorge [Aug.] (1; CUB),
Long Lake ([3140 m]) [July, Sep.] (6; CArm), Mitchell Lake
([3290 m]) [July] (7; CArm, CUB), Rainbow Lakes (10 miles
SW of Ward [3350 m]) [Aug.] (1 ; CNC), Red Rock Lake ([2900
m]) [Aug.] (6; CArm), South St. Vrain Creek ([3050 m]) [July]
(1; CUB), Yankee Doodle Lake ([3140 m]) [July] (1; CArm);
Clear Creek County, Leavenworth Valley (Argentine Road
[2740-3350 m], Waldorf Mine [3540 m]) [June-July] (20; ANSP,
CAS, CNC, DEUN, FMNH, MCZ, ROM, UMMZ, USNM,
ZMLS), Mount Evans (Echo Lake [3230 m], Summit Lake
[3960 m]) [July] (3; CAS, CNC), Silver Plume ([2740-3050
m]) [June] (6; ANSP, CAS, MCZ, UWEM); El Paso County,
Gold Camp Road (9 miles W of Broadmoor at South Cheyenne
Creek [2650-2740 m]) [July-Aug.] (50; CAS, EAMa); Gilpin
County, Rollinsville area [July] (1; CArm); Jackson County,
Cameron Pass ([3 140 m]) [Aug.] (1; UASM); Larimer County,
Bennett Creek [May-July] (18; RTBe), Browns Lake Trail [Aug.]
(1; RTBe), Buckhorn Creek [July] (2; RTBe), Cameron Pass

([3050 m]) [June, Aug.] (6; MSU, RTBe, UASM, USNM),
Crown Point Road (at Bennett Springs [2290-2350 m], at Crown
Point Trail [3140-3200 m], 40 miles W of BeHevue [2740 m])
[June-Aug.] (20; CAS, RTBe, UASM, USNM), Monument
Gulch [July] (18; RTBe), North Fork Cache la Poudre River
[June] (2; RTBe), Zimmerman Lake [Aug.] (5; RTBe), West
Fork Sheep Creek (16 miles W of Teds Place [3050 m]) [Aug.]
(3; CNC); Park County, Kenosha Pass [July] (1; CAS); Rocky
Mountain National Park, Blue Lake [July] (3; RTBe), Chasm
Lake (stream below) [Aug.] (3; CArm), Endovalley Camp-
ground [Aug.] (4; RTBe), Fall River ([2620 m]) [Aug.] (1;
UASM), Hang Lake [Aug.] (1; RTBe), Lake Hiayaha [Aug.]
(3; UAFA), Longs Peak ([3050-3350 m] and Boulder Field
[3840 m], Larkspur Creek) [July-Aug.] (27; CArm, CAS),
Thunder Lake ([3080 m] and Thunder Lake Trail [2500-3350
m]) [June] (2; CArm); Routt County, Walton Creek (above
Dumont Lake [2900-2960 m]) [Aug.] (3; CAS); Teller County,
Divide (8 miles S on Cripple Creek Road [2900 m]) [July] (2;
CNC). WYOMING: Albany County, Brooklyn Lane ([3200
m]) [July] (18; CAS), Centennial [Aug.] (1; MSU), Douglas
Creek (1 mile SSE of Keystone [2440 m]) [July] (16; CAS),
Laramie Peak (south slope at Friend Creek [2440 m]) [July]
(20; CAS), Little Brooklyn Lake ([3120 m]) [July] (8; CAS),
Snowy Range Pass ([3200 m]) [June] (8; UASM); Carbon
County, South Brush Creek (20 miles SE of Saratoga [2470
m]) [July] (1; CAS), Hidden Treasure Gulch (1 1.5 miles WSW
of Encampment [2870 m]) [July] (2; CAS), Silver Lake ([3170
m]) [July] (2; USNM), Slaughterhouse Gulch (1 1 miles SW of
Encampment [2870 m]) [July] (6; CAS).

Specimens Without Locality Data: (1; USNM).
NOMENCLATURAL CHANGES

Since my last two reports on Nearctic Nebria
(Kavanaugh 1979 and 1981), additional speci-
mens and data about geographical and habitat
distribution have also accumulated for previ-
ously described taxa. Study of these specimens
and data, re-examination of some previously
studied materials, and re-evaluation of species
and subspecies concepts developed in previous
reports (Kavanaugh 1978, 1979, and 1981) lead
me to propose the nomenclatural changes pre-
sented below. More complete discussions of pro-
posed synomymies and changes in status of names
will be presented in a forthcoming monography
of Nearctic Nebria.

Nebria intermedia VAN DYKE, 1 949:49 [= Nebria
crassicornis intermedia Van Dyke — NEW
STATUS].

Analysis of the pattern of geographical varia-
tion shown by samples representing Nebria cras-
sicornis Van Dyke (1925:121) and N. intermedia
Van Dyke throughout their ranges suggests that
these taxa represent allopatric forms that are
clearly differentiated but not to a degree typical
of closely related, sympatric species. I suggest

KAVANAUGH: NEW NEARCTIC NEBRIA

167

that they be considered subspecies of a single
species, N. crassicornis.

Nebria sonorae KAVANAUGH, 1981:438 [=Ne-
bria acuta sonorae Kavanaugh— NEW STATUS].

My description of N. sonorae was based on
only one male and two female specimens. These
specimens were only slightly, but consistently,
different from specimens of Nebria acuta acuta
Lindroth in several characters of external struc-
ture. However, form of the aedeagus of the male
specimen (chosen as holotype) was so different
from that of TV. acuta acuta males that I had no
doubt concerning specific distinctiveness of the
two taxa.

Several additional male specimens of N. so-
norae recently collected and studied all have ae-
deagi with form typical of N. acuta acuta males.
I now conclude that the aedeagus of the holotype
male of TV. sonorae is not typical for that taxon
and, further, that the two forms are conspecific.
However, differences in external structural char-
acters between specimens of TV. sonorae and TV.
acuta acuta are both consistent and sufficient to
support their status as allopatric subspecies of a
single species.

Nebria fragilis CASEY, 1924:21 [=Nebria arkan-

sanafragilis Casey— NEW STATUS].
Nebria arkansana uinta Kavanaugh, 1979:102

[= Nebria arkansana fragilis Casey — NEW

SYNONYMY].
Nebria fragilis teewinot Kavanaugh, 1979:103

[= Nebria arkansana fragilis Casey — NEW

SYNONYMY].

Males of Nebria arkansana arkansana Casey
and TV. a. edwardsi Kavanaugh differ distinctly
from males of "Nebria fragilis Casey" in form
of aedeagus and in several characters of external
structure. Male specimens from the Uinta and
northern Wasatch Mountains of northern Utah
share aedeagal form with males of the two TV.
arkansana subspecies just mentioned, yet differ
from them slightly in characters of external struc-
ture. I therefore recognized (Kavanaugh 1979)
these specimens as representing a distinct sub-
species of TV. arkansana, namely TV. a. uinta. Male
specimens from the Teton, Wind River, and ad-
jacent Mountains in western Wyoming share ae-
deagal form with males of N. fragilis; but, again,
they differ from the latter in several characters
of external structure and color. Based on these

similarities and differences, I recognized (Ka-
vanaugh 1979) the Wyoming specimens as rep-
resenting a distinct subspecies of TV. fragilis,
namely TV. / teewinot. The resulting pattern of
geographical distribution was one in which the
range of all subspecies of both TV. arkansana and
TV. fragilis were mutually allopatric, although
ranges of TV. arkansana uinta and TV. fragilis fra-
gilis were essentially parapatric in northcentral
Utah.

Since 1979 I have studied additional material
from areas that previously represented gaps be-
tween the allopatric ranges of described subspe-
cies, and I re-examined specimens studied ear-
lier. These studies have shown that samples of
males from localities in presumed gaps, as well
as some samples from localities bordering these
gaps, are mixed in aedeagal form and interme-
diate in characters of external structure in rela-
tion to respective allopatric forms. Such findings
lead me to conclude that TV. arkansana and TV.
fragilis are conspecific. Specimens representing
TV. arkansana uinta, N. fragilis fragilis, and TV./
teewinot all share a combination of structural
features which distinguish them from members
of other TV. arkansana subspecies. I therefore
suggest that these forms (and respective inter-
mediates between them) together represent a sin-
gle, distinct subspecies of TV. arkansana for which
the name TV. arkansana fragilis has priority. A
reconstruction of the historical development of
the complex pattern of geographical variation
within this subspecies will be presented in the
monograph now in preparation.

Nebria tr if aria tetonensis ERWIN AND BALL, 1 972:
95 [=Nebria trifaria trifaria LeConte— NEW
SYNONYMY].

Based on an almost continuous series of sam-
ples (all collected after 1972) from the area be-
tween respective type localities for TV. trifaria
trifaria and TV. trifaria tetonensis, I suggest that
these two nominal taxa represent simply the ex-
tremes of continuous clinal variation in those
characters previously used to distinguish their
members.

Nebria trifaria piute ERWIN AND BALL, 1972:95
[=Nebria piute piute Erwin and Ball— NEW
STATUS].

Members of Nebria piute and TV. trifaria
LeConte differ from each other at least as much

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12

as other closely related, but sympatric, species
in characters of both external structure and gen-
italia (of both males and females). I therefore
suggest that they represent distinct species.

Nebria trifaria utahensis KAVANAUGH, 1979: 110
[=Nebria piute utahensis Kavanaugh— NEW
STATUS].

Based on form of aedeagus of males and bursa
copulatrix of females, TV. utahensis, described as
a subspecies of N. trifaria LeConte, should in-
stead be considered a subspecies of N. piute, along
with N. piute sevieri n.ssp. described above.

LITERATURE CITED

CASEY, T. L. 1924. Additions to the known Coleoptera of

North America. Memoirs on the Coleoptera, 1 1:1-347.
ERWIN, T. L., AND G. E. BALL. 1972. Classification of the

ovipennis and trifaria groups of Nebria Latreille (Coleoptera:
Carabidae: Nebriini). Proceedings of the Biological Society
of Washington, 85:77-108.

KAVANAUGH, D. H. 1978. The Nearctic species of Nebria
Latreille (Coleoptera: Carabidae: Nebriini): classification,
phylogeny, zoogeography, and natural history. Unpublished
Ph.D. dissertation. Department of Entomology, University
of Alberta, xlviii +1041 pp.

. 1979. Studies on the Nebriini (Coleoptera: Carabi-
dae), III. New Nearctic species and subspecies, nomencla-
tural notes, and lectotype designations. Proceedings of the
California Academy of Sciences, 42:87-133.

. 1981. Studies on the Nebriini (Coleoptera: Carabi-

dae), IV. Four new Nebria taxa from western North America.

Proceedings of the California Academy of Sciences, 42:435-

442.
VAN DYKE, E. C. 1925. Studies of western North American

Carabinae with descriptions of new species. The Pan-Pacific

Entomologist, 1:111-125.
. 1949. New species of North American Coleoptera.

The Pan-Pacific Entomologist, 25:49-56.

KAVANAUGH: NEW NEARCTIC NEBRIA

169

FIGURES 1-8. Fig. 1. Right antennal scape, dorsal aspect, Nebria gebleri albimontis n.ssp. (Birch Creek, California); scale
line = 1.0 mm. Figs. 2-8. Pronotum, dorsal aspect; scale line = 1.0 mm. 2. Nebria altisierrae n.sp. (Olmsted Point, California).
3. Nebria campbelli n.sp. (Mount Baker, Washington). 4. Nebria wallowae n.sp. (West Fork Wallowa River, Oregon). 5. Nebria
jeffreyi n.sp. (South Fork McCoy Creek, Oregon). 6. Nebria haida n.sp. (Mount Needham, Queen Charlotte Islands, British
Columbia). 7. Nebria louiseae n.sp. (Skedans, Queen Charlotte Islands, British Columbia). 8. Nebria gebleri albimontis n.ssp.
(Birch Creek, California).

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12

FIGURES 9-14. Pronotum, dorsal aspect; scale line = 1.0 mm. 9. Nebria labontei n.sp. (West Fork Wallowa River, Oregon).
10. Nebria calva n.sp. (Mount Baldy, Arizona). 1 1. Nebria sierrablancae n.sp. (Sierra Blanca, New Mexico). 12. Nebria piute
sevieri n.ssp. (Parowan Creek, Utah). 13. Nebria steensensis n.sp. (South Fork McCoy Creek, Oregon). 14. Nebria trifaria
pasquineli n.ssp. (Lefthand Creek, Colorado).

KAVANAUGH: NEW NEARCTIC NEBRIA

171

FIGURES 15-16. Basal region of left elytron, dorsal aspect; scale line = 1.0 mm. 15. Nebria altisierrae n.sp. (Olmsted Point,
California). 16. Nebria campbelli n.sp. (Mount Baker, Washington).

FIGURES 1 7-20. Median lobe of male genitalia, left lateral aspect (or a = left lateral aspect, b = ventral aspect); scale line =
1.0 mm. 17. Nebria wallowae n.sp. (West Fork Wallowa River, Oregon). 18. Nebria gebleri albimontis n.ssp. (Birch Creek,
California). 19. Nebria calva n.sp. (Mount Baldy, Arizona). 20. Nebria sierrablancae n.sp. (Sierra Blanca, New Mexico).

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12

FIGURES 21-23. Median lobe of male genitalia (a = left lateral aspect, b = ventral aspect, c = apical aspect); scale line = 1.0
mm. 21. Nebria piute sevieri n.ssp. (Parowan Creek, Utah). 22. Nebria steensensis n.sp. (South Fork McCoy Creek, Oregon).
23. Nebria trifaria pasquineli n.ssp. (Lefthand Creek, Colorado).

KAVANAUGH: NEW NEARCTIC NEBRIA

173

FIGURES 24-26. Bursa copulatrix of female (a. dorsal aspect; b. left lateral aspect; c. mid-sagittal outline, left lateral aspect);
scale line = 1.0 mm. 24. Nebria wallowae n.sp. (West Fork Wallowa River, Oregon). 25. Nebria gebleri albimontis n.ssp. (Birch
Creek, California). 26. Nebria calva n.sp. (Mount Baldy, Arizona).

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12

FIGURES 27-28. Bursa copulatrix of female (a. dorsal aspect; b. left lateral aspect; c. mid-sagittal outline, left lateral aspect);
scale line = 1.0 mm. 27. Nebria sierrablancae n.sp. (Sierra Blanca, New Mexico). 28. Nebria piute sevieri n.ssp. (Parowan Creek,
Utah).

KAVANAUGH: NEW NEARCTIC NEBRIA

175

FIGURES 29-30. Bursa copulatrix of female (a. dorsal aspect; b. left lateral aspect; c. mid-sagittal outline, left lateral aspect);
scale line = 1.0 mm. 29. Nebria steensensis n.sp. (South Fork McCoy Creek, Oregon). 30. Nebria trifaria pasquineli n.ssp.
(Lefthand Creek, Colorado).

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 12

4 0

FIGURE 3 1 . Map of geographical distributions: Nebria altisierrae n.sp. = solid circle; Nebria campbelli n.sp. = solid diamond;
Nebria wallowae n.sp. = solid square; Nebria jeffreyi n.sp. = open diamond; Nebria haida n.sp. = solid triangle; Nebria louiseae
n.sp. = open triangle.

KAVANAUGH: NEW NEARCTIC NEBRIA

177

FIGURE 32. Map of geographical distributions: Nebria gebleri albimontis n.ssp. = solid square; Nebria labontei n.sp. = solid
triangle; Nebria calva n.sp. = solid diamond; Nebria sierrablancae n.sp. = open triangle; Nebria piute sevieri n.ssp. = inverted
solid triangle; Nebria steensensis n.sp. = open diamond; Nebria trifaria pasquineli n.ssp. = solid circle.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 13, pp. 179-220, 22 figs., 2 tables. July 12, 1984

SKELETAL ANATOMY AND CLASSIFICATION

OF THE NEOTENIC ASIAN SALMONIFORM SUPERFAMILY

SALANGOIDEA (ICEFISHES OR NOODLEFISHES)

By
Tyson R. Roberts

California Academy of Sciences, Golden Gate Park, San Francisco, California 94118

ABSTRACT: The distinctive, largely cartilaginous skeletal anatomy of the Asian icefishes or noodlefishes is
described and figured from cleared specimens stained with alcian and alizarin. This information, together
with examination of types and other material in North American and European museums, leads to the
following revised classification of these neotenic Salmoniformes:
Salangoidea new superfamily (coordinate with Osmeroidea, Salmonoidea)

Salangidae

Protosalanginae: Protosalanx chinensis

Salanginae: Salanx (Salanx) ariakensis, S. (S.) cuvieri, S. (Hemisalanx) prognathus, S. (Leucosoma) reevesi

Salangichthyinae new subfamily: Neosalanx andersoni, N, brevirostris, N. Jordan!, N. reganius, Salang-
ichthys ishikawae, S. microdon

Sundasalangidae: Sundasalanx microps, S. praecox

The introduction includes a summary of salangoid natural history and a key for their identification. The
systematic account includes all primary and secondary synonyms of genera and species recognized. New
information is presented on pectoral girdle morphology in teleosts, relationships of salangoids and other
salmoniforms, and breeding tubercles, meristic variation, and neoteny in salangoids.

INTRODUCTION Japanese, however, they are usually referred to

The slender, soft-bodied, and transparent or as shirauwo (whitefishes) and rarely as hiagio

translucent salmoniform fishes of the family Sa- (icefishes). In Russian they are usually referred

langidae inhabit the sea coasts, rivers, and lakes to as lapsha-ryba or noodlefish, and an equiva-

of East Asia including Japan from Sakhalin, Via- lent name exists in Chinese, mien-tiao-yu

divostok, and the Amur River south to northern < *** >• They have been referred to as Nudel~

Vietnam (Tonkin). The greatest concentration of ^^ . in Ge™an'. but noodlefishes> a "* aP'

, ...._. - propnate and distinctive name, seems not to have

genera and species is in China and Korea. Of 1 1 , . „ ,. , , ^ .,

. " • appeared in English except m a translation of a

species herein recognized, eight occur in China, Russian wQrk (Berg { %2.480) Thg fle§h i§ tasty5

eight or nine m Korea, and four m Japan. Only whether cooked as a SQU^ eaten whh vin£gar Qr

Salangichthys microdon occurs along the outer scrambled eggs, or fried (Okada 1955:60). The

coast of Korea and in Siberia, and only Salanx species most commonly eaten in Japan is Sa-

reevesi and Neosalanx brevirostris have been re- langichthys microdon, and in China probably

ported as far south as Tonkin (or Haiphong). Neosalanx brevirostris or N. jordani. Protosa-

Members of the Salangidae have almost al- lanx and Salanx are also consumed, but I doubt

ways been referred to in English as icefishes. In that tiny Sundasalanx has ever been dined upon.

[179]

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

FIGURE 1 . Axial skeleton, (a) Protosalanx chinensis, CAS-SU 6306, 85.5-mm adult male; (b) Salanx cuvieri, CAS-SU 32454,
6 1.7-mm juvenile sex undetermined; (c) Salangichthys ishikawae, CAS 6780, 74-mm adult female; (d) Neosalanx jordani, CAS
52028, 38.3-mm adult male; (e) Sundasalanx microps, CAS 44220, 17-mm adult sex undetermined.

Despite their standing as a delicacy— sufficient
for them to be imported by the Chinese and Jap-
anese communities of San Francisco and served
in the city's sushi bars— relatively little is known
about the systematics and biology of noodlefish-
es. An impression of their morphological diver-
sity can be obtained from Figures 1 and 2.

The present study was undertaken in connec-
tion with the discovery of some minute, scaleless,
and transparent fishes during my fieldwork in the

Malay Peninsula (1 97 1 , 1973) and on the Kapuas
River in Kalimantan Barat, Indonesia (1976).
When first found, although in fresh water, they
were living close to the sea and were mistaken
for elopoid leptocephali, which they resemble
only superficially. In the Kapuas River, however,
they were living 800 km upriver in the midst of
a rich riverine fish fauna dominated by Ostar-
iophysi and with no elopoids. The observation
that the maxillary bones curved inwards below

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

181

TABLE 1. REPRODUCTIVE BIOLOGY OF SALANGOIDS (Wakiya and Takahasi 1937, Okada 1960, Senta 1973a, b, Roberts 1981,
and pers. obs.).

Larg-

Smallest

Larg-

est

Diam-

mature

est

fe-

eter

male

egg

(mm)

Fecundity Remarks

Protosalanx chinensis

168

146

1.15

Breeds January-February (Korea)

Salanx ariakensis

—

147

0.75

Breeds October-November (Korea)

Salanx cuvieri

—

144

—

— —

Salanx prognathus

100

Ill

119

0.85

Breeds April-May

Salanx chinensis

130

153

—

— —

Neosalanx andersoni

100

—

Breeds April-May (Korea)

Neosalanx brevirostris

—

0.7

— —

Neosalanx jordani

56.5

59.5

0.5

Breeds March-May

Neosalanx reganius

—

0.9

Breeds February-March

Salangichthys ishikawae

—

0.95

Breeds April-May

Salangichthys microdon

100

0.91-0.99

1300-2700 Breeds March-May

Sundasalanx microps

—

Largest specimen (sex unknown)

19.9 mm

Sundasalanx praecox

14.9

18.3

17.3

0.20-0.25

50 Both sexes ripe in June

the head led to an hypothesis that they are sa-
langoids, and observations of their skeletal anat-
omy and particularly the suspensorium con-
firmed this (Roberts 198 1). These fishes differ in
a number of respects from Salangidae and con-
stitute a separate family, Sundasalangidae, with
one genus, and two or more species, one in the
Malay Peninsula and one or two in the Kapuas
River (Roberts 1981). Sundasalanx also occur
in the Mekong basin, as reported herein. This is
the only truly tropical genus in the entire order
Salmoniformes. Sundasalanx praecox, with
males and females sexually ripe at only 1 4.9 mm,
is the smallest member of the order, and provides
a striking example of a minute secondary fresh-
water fish living in the midst of a rich freshwater
ichthyofauna dominated by primary freshwater
Ostariophysi.

Interest in Sundasalangidae and its relation-
ships led me to examine other salangoids but my
observations and drawings quickly became too
extensive to incorporate in the original descrip-
tion of the new taxa; hence the present mono-
graph.

Food Habits

All salangoids, including tiny Sundasalanx,
appear to be predators. The largest species, Pro-
tosalanx chinensis and Salanx reevesi, both with
well-developed teeth on the tongue and jaws, ap-
parently feed mainly on fishes. Salangichthys

microdon taken in the Takahashi River had fed
on larvae of the goby Chaenogobius sp. and on
the mysid shrimp Neomysis sp. (Senta 1973b).
Other species of Salanginae and Salangichthyi-
nae feed mainly on small Crustacea (in marine
environments) or on insects (in fresh water).
Sundasalanx are known only from fresh water
and feed on tiny insects (Roberts 1981).

Reproduction

While some species are primarily marine or at
least brackish water inhabitants (e.g., Protosa-
lanx chinensis), and many spend part of their
lives in the sea, others are restricted to fresh water
or have populations which presumably repeat
their life cycle without leaving fresh water. Basic
information on salangoid reproductive biology
is summarized in Table 1 . Fecundity ranges from
several thousand eggs in Protosalanginae and
Salanginae (no precise numbers available) down
to only about 50 in Sundasalangidae.

The external egg membrane is adhesive, eggs
becoming attached to any solid object at the
spawning site. Wakiya and Takahasi (1937, pi.
21) published drawings of the basal portion of
the adhesive strands on the eggs of Protosalanx
chinensis, Salanx ariakensis and S. prognathus,
Salangichthys microdon and S. ishikawae, and
Neosalanx jordani. The eggs illustrated are pre-
sumably ovarian, since the adhesive strands are
not detached. For photomicrographs of the

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

FIGURE 2. Radiographs, (a) Salanx cuvieri, MNHN 9900, 1 12 mm, holotype; (b) Neosalanx andersoni, NRM 10287, 79
mm, holotype.

spawned eggs with detached adhesive threads of
unfertilized and fertilized eggs of Salangichthys
microdon see Okada (1960, pi. 18). Spawning
ecology of this species is described by Senta
(1973a). According to Wakiya and Takahasi
(1937:269), after spawning "the body becomes
very lean and the vertebrae become visible
through the skin, whence it is generally assumed
that death then ensues." I suspect that this is true
in Salanginae as well as Salangichthyinae but not
in Protosalanx.

Sexual Dimorphism

A notable feature of salangoids is their unique
sexual dimorphism. In all Salangidae except
Neosalanx, sexually mature males have the pec-
toral fins longer and more pointed (falcate) and
the pelvic fins larger. In all adult male Salangidae
the anal fin is larger than in females and has
modified rays. The anterior rays of the anal fin
are greatly enlarged, the middle rays thin and
strongly curved, and the posterior rays short and
widely separated at the base. The morphology of
the anal fin is very similar in sexually mature
males of all of the genera and species of Salan-
gidae. In all Salangidae, mature males have a row
of large, tightly adherent scales on the body par-
allel to the anal fin base (sometimes extending
posteriorly a short distance beyond the anal fin
base onto the caudal peduncle). The number of
anal scales ranges from 14 to 28. Sexual dimor-

phism has not been observed in Sundasalangi-
dae.

Although salangids differ greatly in the size of
adult males, the morphology of the modified male
anal fin is remarkably uniform (Fig. la, d). The
total range of anal fin-rays is 23-32. The first two
or three rays are simple, the first one or two small
or minute. The last simple ray and the first four
to six branched rays are greatly enlarged and
somewhat thickened; near the base of each of
these rays is a very large lateral projection. The
next 1 2 or so rays are noticeably thinner and are
deflected backwards near the middle of their
length, so that their distal portions lie close to-
gether. In Protosalanx these rays are simple, but
in other Salangidae they are branched. The pos-
teriormost rays may be simple or branched, are
reduced in size and not modified, except that
their bases tend to be relatively wide apart (much
more so than the bases of the preceding rays or
of the corresponding rays in females), especially
in Salanginae. The proximal pterygiophores, es-
pecially for the anterior portion of the anal fin,
are also enlarged in males. In alcian-alizarin
preparations the anal fin-rays and pterygiophores
of sexually mature males are deeply stained with
alizarin, whereas those of females tend to be less
well stained with alizarin or in some instances
stained only with alcian.

Near the middle of the rays in the most mod-
ified part of the male anal fin, a tough, almost
tendonlike membrane arises from each ray and

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

183

extends obliquely and posteroventrally across the
densely webbed portion of the fin to end in a
thickened, obliquely oriented non-muscular pad
of tissue. The distal portion of this oblique pad
is free from the surface, so it can be readily lifted,
and even when not lifted forms a sort of groove
for the length of the pad. This portion of the anal
fin can be flexed in such a way that it forms a
slight concavity. The fin may be expanded man-
ually by pulling on the anteriormost rays; when
released, it snaps back into a less expanded con-
dition. Spawning behavior has not been reported
upon, but presumably the male's anal fin remains
in contact with the vent region of the female in
such a way that it temporarily retains eggs and
sperm in proximity while fertilization occurs ex-
ternally.

In addition to the modified anal fin, sexually
mature males of all Salangidae bear a row of
large, cycloid scales on the side of the body above
and co-extensive with the anal fin or extending
a short distance beyond it onto the caudal pe-
duncle. The scales are tightly adherent and
broadly overlapping (more so anteriorly than
posteriorly). In addition to the main row of anal
scales, some specimens exhibit two or three
smaller scales in a separate row overlying the
vent. These usually have been overlooked by
previous authors, and are not included in the
counts of anal scales in Table 2.

Breeding tubercles and other forms of tem-
porary sexual dimorphism have not been re-
ported previously in salangoids. I have observed
breeding tubercles in adult males and females,
apparently in spawning condition, of Protosa-
lanx chinensis, and in adult males of Salangich-
thys microdon and Neosalanx jordani. This pre-
sumably temporary tuberculation is most
extensive and easily observable in an 120-mm
male Protosalanx (CAS-SU 36025). In this spec-
imen breeding tubercles occur on the anal, pec-
toral and pelvic fins, abdominal keel, and head.
The strong lateral projections on the anterior face
of the first nine branched anal fin-rays are en-
tirely or almost entirely covered by a thickened,
longitudinal band of thickened skin 9 mm long
and 1.2 mm high. The surface of this spongy
band of skin is covered with hundreds of small,
overlapping, scale- or leaflike breeding tuber-
cles, with their raised free margins projecting an-
teriorly. There are about 1 2-20 of these tubercles
in a vertical series. Discrete pads of similarly

thickened skin covered with similar breeding tu-
bercles extend obliquely posteroventrally on the
basal third of the first five branched anal fin-rays.
There are up to eight tubercles across each ray.
The skin on the middle third of the same rays
appears to be only slightly thickened and bears
only a few, small widely spaced, low-lying round
(not scalelike) tubercles. The distal third or
branched portion of the first eight branched rays
is covered with thick skin densely coated with
scalelike tubercles. There are up to about eight
tubercles across each ray-branch. The leading edge
of the third (enlarged) simple anal fin-ray bears
a thick, lamellar projection of skin, 11.5 mm long
and up to 2.2 mm wide, covered with widely
scattered, low-lying round tubercles without free
margins. The midventral abdominal keel is also
notably thickened, and covered with minute,
closely spaced round or granular tubercles which
extend for a short distance onto the abdomen
and sides of the body just anterior to the anal
fin. The pelvic and pectoral fins bear round tu-
bercles dorsally and ventrally; these are most no-
ticeable on the enlarged outermost pectoral fin-
ray. The dorsal fin is slightly tuberculate, the
adipose and caudal fins non-tuberculate. The
dorsal, lateral, and ventral surfaces of the head
bear irregularly scattered, round, low-lying tu-
bercles without free margins. These are largest
and most numerous on its ventral surface. The
skin of the oral margin of the upper and lower
jaws and gular margin of the lower jaw is thick-
ened and tuberculate. Fine granular projections,
which may be minute breeding tubercles, extend
in a dorsomedian longitudinal band from the
dorsal fin origin anteriorly halfway to the occi-
put. In the two gravid females the skin is less
modified, and although tuberculation is very
much lighter, there are small, low-lying round
tubercles on the anal, pelvic, and pectoral fins
and on the head. In one of them the skin on the
jaws is thickened as in the male; in the other it
is not. The first female has the median abdominal
fold somewhat thickened, suggestive of the more
pronounced thickening of this fold seen in the
male; the other female does not. Tubercles have
not been observed in females of any other sa-
langoid.

In other salangoids breeding tubercles have
been observed only on the anal fin of males. An
8 3.1 -mm male Salangichthys microdon (CAS
52033) has small scalelike breeding tubercles on

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

the branched portion of the anteriormost
branched anal fin-rays. These are arranged uni-
serially on each fin-ray branch. A 47.5-mm male
Neosalanx jordani (AMNH 51704) has very
similar scalelike tubercles on thickened skin sur-
rounding the lateral projection at the base and
on the basal half of the first four branched anal
fin-rays (which are enlarged); fin-rays 6-14, which
are bent, each have three to four melanophores
on the basal one-fourth of their length. (Similar
coloration has been observed on the anal fin in
occasional males of Salangichthys microdon.)

Breeding tubercles apparently do not occur in
Sundasalangidae, in which neither secondary
sexual dimorphism nor dichromatism has been
observed.

Pigmentation

The only pigmentation known to be exhibited
by salangoids, apart from that of their eyes, is in
melanocytes or melanophores, which tend to oc-
cur as widely separated single cells or isolated
clumps of relatively few cells. In life all, or almost
all, salangoids (except Protosalanx) are trans-
parent or translucent, except for the prominent
eyes. The most constant pigmentary feature of
the salangoids is a row of melanophores at the
interface of the ventral myotomic musculature
and the non-segmentally muscularized ventral
abdominal wall. This series of melanophores,
with a single cell at about the middle of the ven-
tral end of each myotome, from the most anterior
myotome to the anal fin origin, is present in near-
ly all salangoid specimens examined. Usually
these melanophores are longitudinally elongate,
giving the appearance of a series of widely spaced
thin black dashes. A second pigmentary feature
found in many salangoids is a ventromedian row
of widely spaced melanophores, one for each body
segment. These melanophores tend to be den-
dritic when expanded or round when contracted,
and may extend the entire length of the abdomen;
sometimes they are restricted to the preanal
membranous keel. These two pigmentary fea-
tures of salangoids occur in many teleost larvae
and in adults of other neotenic teleosts.

Some salangoids exhibit a row of melano-
phores along the anal fin base, one between each
anal fin-ray. This row of melanophores, lying
deep in the body and median rather than paired,
may be the continuation of the midabdominal

row of melanophores described above. This row
usually extends the length of the anal fin; some-
times it continues beyond the anal fin onto the
caudal peduncle near its ventral margin.

Clusters of a few melanophores occur just an-
terior to the bases of the pectoral and pelvic fins
in most salangoids, at the tip of the snout and
chin, especially in Salangichthys, and infre-
quently on the dorsal surface of the head over-
lying the fore- and hind-brain. In sexually mature
(spawning?) males of Salanx and Salangichthys
there may be a cluster of melanophores on the
proximal portion of the middlemost anal fin-
rays. The dorsal, anal, pectoral, and pelvic fins
are otherwise usually devoid of melanophores,
but the caudal fin lobes frequently are dark or
dusky due to numerous fine melanophores. The
anal scales of the males are always entirely de-
void of melanophores.

In most salangoids the entire dorsal and most
of the lateral body surfaces are devoid of mela-
nophores. Protosalanx chinensis and Neosalanx
andersoni provide notable exceptions. Young of
Protosalanx and. Neosalanx exhibit very few me-
lanophores. Large and sexually ripe individuals
of these two species, however, may have the dor-
sal and lateral surfaces of the body with numer-
ous melanophores. Those on the dorsal body sur-
face are fine, exceedingly numerous, and generally
scattered over the entire musculature, but those
on the sides are few and peculiarly restricted along
the course of the myotomal septae. About a doz-
en melanophores lie on each myotomal septa;
the melanophores of successive septae are more
or less parallel to each other; the cells are oblique-
ly elongate, conforming to the thinness and
obliquity of the septae and thus forming a series
of widely spaced thin black slashes. This pattern,
sometimes barely evident or absent in P. chi-
nensis, is very well developed in two gravid fe-
males of 1 29-132 mm (USNM 1 20746). Wakiya
and Takahasi (1937) show it well developed in
female P. chinensis (not gravid?) and N. ander-
soni (gravid); and relatively weakly developed in
males of both species. It is present only on the
upper part of the body in the relatively small
male holotype of N. andersoni (NRM 10287, 79
mm). Chyung (1961) shows it well developed in
a gravid N. andersoni. I have seen clupeomorphs
but no osmeroids or other salmoniforms with
similarly distributed melanophores.

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

185

Some Misconceptions

Some misconceptions about Salangidae should
be noted. These concern the reported presence
of scales other than anal scales, presumed ab-
sence of the swimbladder, and reputed oral
brooding of eggs. The most persistent misinfor-
mation concerns the occurrence of scales on the
body other than the anal scales. Gunther (1866:
205) stated that the body is "naked or covered
with small, exceedingly fine, deciduous scales (?)"
and added in a footnote, "There is no trace of
scales in specimens preserved in spirits for some
time; but others, which we received lately, show
scattered fragments of scales, without any regular
arrangement." He was unaware of the anal scales
of males. Regan (1908b:444), in diagnosing Sa-
langinae (= Salangidae), stated simply "scales de-
ciduous" but described the anal scales of males
in a footnote. Fang ( 1 934a:239) stated body "na-
ked or with a few exceedingly thin, large, scat-
tered, deciduous scales, without any regular ar-
rangement" in addition to the anal scales of males.
Nichols (1944) referred to several species with
"scales small, deciduous, little evident." Nelson
(1976:104) cautiously stated "body generally
scaleless" without referring to the anal scales of
males. As noted by Wakiya and Takahasi (1937)
all salangids are totally scaleless except for the
anal scales of sexually mature males; as noted
above, the anal scales are large and strongly ad-
herent. Reports of scales on other parts of the
body are all attributable to dislodged scales from
other fishes.

Various authors, including Gunther (1866:
205), Fang (1934a:239), and Nelson (1976:104)
have stated that salangids lack a swimbladder.
Wakiya and Takahasi (1937:268, fig. 1) reported
a physostomous swimbladder in Protosalanx
chinensis, Salanx ariakensis, S. prognathus,
Neosalanx jordani, Salangichthys ishikawae, and
S. microdon. In P. chinensis and S. ishikawae
the swimbladder is depicted as relatively large
and oval, and in the others as equally long but
almost uniformly slender for its entire length.
The condition of the swimbladder in Sundasa-
langidae is unknown.

Fang (1934a:238, 252, fig. 7) suggested that
Salangidae are oral brooders. In a series of 6 1
males and 27 females identified as Hemisalanx
(=Salanx) prornathus collected at Chinkiang in

April 1933, Fang found 6 males and 19 females
with 1-21 eggs in the mouth. He also reported
one Protosalanx (sex not mentioned) with eggs
in its mouth. I have also observed a few speci-
mens of both sexes, especially of Salanginae, with
small numbers of eggs in the mouth; this is at-
tributable to rupture of the ovaries and spillage
of eggs after the fish had been caught. There is
no information indicating that salangoids prac-
tice oral brooding or any other form of parental
care.

This introduction to salangoids concludes with
a key for their identification.

Key to Salangoidea

1 a. Pelvic fin with 5 rays; adipose fin absent;
pectoral fin rayless throughout life; sex-
ually mature males without anal scales
or enlarged anal fin; vertebrae 37-43;
standard length to 22 mm (Sundasalan-
gidae) 11

1 b. Pelvic fin usually with 7 rays (rarely 6 or
8); adipose fin present; pectoral fin with
rays except in larvae; sexually mature
males with a row of large anal scales and
enlarged anal fin; vertebrae 48-79; adults
at least 35 mm in standard length (Sa-
langidae) 2

2a. Teeth on palatal toothplate and lower
jaw in two rows; teeth on tongue in two
marginal rows or widely spread over ba-

sihyal toothplate (Protosalanginae)

Protosala nx ch inensis

2b. All oral teeth in single rows 3

3a. Head extremely depressed; snout very
elongate and relatively pointed; cranial
fontanel entirely closed in juveniles and
adults; premaxillae larger than maxillae,
those of opposite sides meeting broadly
in front of snout; premaxillary teeth rel-
atively large; supramaxilla absent; ver-
tebrae 66-79 (Salanginae) 4

3b. Head moderately depressed; snout mod-
erately elongate and broadly rounded;
cranial fontanel with anterior and pos-
terior portions open throughout life, pre-
maxillae smaller than maxillae, more or
less separated from each other in front
of snout; premaxillary teeth relatively
small, tiny, or absent; supramaxilla pres-

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

ent; vertebrae 48-65 (Salangichthyinae)

4a. Tongue with a median row of conical

teeth (subgenus Leucosomd)

Salanx reevesi

4b. Tongue toothless 5

5a. Head strongly pointed; lower jaw not
projecting beyond upper jaw; presym-
physeal fleshy appendage, bone, and teeth
frequently present in adults; vertebrae
72-79 (subgenus Salanx) 6

5b. Head less strongly pointed; lower jaw
projecting slightly beyond upper jaw; no
presymphyseal fleshy appendage, bone,
or teeth; vertebrae 70-73 (subgenus
Hemisalanx) Salanx prognathus

total rakers on first gill arch 9-15 (main-
land Asia) Neosalanx jordani

lOc. Vertebrae 52-56, average 53.75 (after
Wakiya and Takahasi 1937); standard
length to 58 mm; total rakers on first gill
arch 15 (known only from Ariake Bay,
Kyushu, Japan) Neosalanx reganius

I la. Horizontal diameter of eye less than 4%

of standard length; ceratobranchial 5 with
0-3 small conical teeth; total rakers on

first gill arch 0-2; vertebrae 41-43

Sundasalanx microps

I 1 b. Horizontal diameter of eye more than

5% of standard length; ceratobranchial 5
with about 8-10 large conical teeth; total
rakers on first gill arch 10-12; vertebrae
37-41 Sundasalanx praecox

6a. Presymphyseal bone usually present in

specimens over 1 00 mm standard length, MATERIAL EXAMINED
relatively elongate and with up to 1 7 teeth Salangoid specimens deposited in the follow-
on each side; vertebrae usually 77-78 ing institutions have been examined for this study:
(rarely 76 or 79) Salanx cuvieri American Museum of Natural History, AMNH:

6b. Presymphyseal bone usually absent, or British Museum (Natural History), BMNH; Cal-
relatively short and with no more than ifornia Academy of Sciences, CAS, including
6 teeth on each side; vertebrae 72-75 ... specimens formerly deposited at Stanford Uni-
Salanx ariakensis versity, CAS-SU; Museum national d'Histoire

_ „ . . , naturelle, Paris, MNHN; Naturhistoriska Riks-

7a. Palatal toothplate with minute teeth; '^ , ' XTr,A(, »„ r~ ,

... f, . museet, Stockholm, NRM: Museum of Zoology,

premaxilla with numerous small or mi- TT . . r»*-il- T™™*^ o • u

A, 1^-11 University of Michigan, UMMZ; Smithsonian

nute teeth, snout relatively elongate; ver- T . . TT0^Tnk, \ ~ , . , »,

cn' - /0 , . ,,, ' o Institution, USNM; and Zoologisch Museum,

tebrae 59-65 (Salangichthys) 8 TT . ' „,?,.

-,, r» , , , , ... „ Umversiteit van Amsterdam, ZMA.

7b. Palatal teeth absent; premaxilla usually . , , .. ,.. , r . . . , ,. , ,.

iU . -.!_ 1 c • .L A detailed list of material examined (including

toothless or with 1-5 minute teeth; snout . . .. ....

. . , , „. . , , , alcian-alizann preparations) is given under each

relatively short except in Neosalanx an- . . .

-o,c/,, , , ~ species in the systematic account.

aersom; vertebrae 48-65 (Neosalanx) 9

8a. Pectoral fin-rays 14-19 SKELETAL ANATOMY

Salangichthys microdon Salangoid skeletal anatomy cannot be ob-

8b. Pectoral fin-rays 20-28 served adequately from alizarin preparations be-

Salangichthys ishikawae cause it is largely cartilaginous, and even ossified

9a. Snout relatively short, standard length to Portions (induding dermal bones) often fail to

64 mm, males with 14-21 anal scales, stam wflth allzann' The only Previous obs^rva-

vertebrae fewer than 60 10 tlons of salangoid skeletal anatomy are brief and

9b. Snout relatively elongate, standard length relatively uninformative. The only general ac-

to 100 mm, males with 20-28 anal scales, count' that of McDowall (1969:815), is limited

vertebrae 63-65 Neosalanx andersoni to three Paragraphs, one on the cranium, one on

the jaws, and one on the remainder of the skel-

lOa. Vertebrae 55-59; standard length to 64 eton emphasizing the median fins. Wakiya and

mm; total rakers on first gill arch 15-19 Takahasi (1937) figured toothed portions of the

(mainland Asia) Neosalanx brevirostris jaws, palate, and tongue of various salangids.

1 Ob. Vertebrae usually 50-53, rarely 49 or 54; Nelson ( 1 970) described and figured the gill arch-
standard length usually less than 50 mm; es in Salanx reevesi and Neosalanx brevirostris

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

187

parietal

dilatator fossa

supraethmoid

frontal

nasal

cranial fontanel

trabecula communis

ethmoid plate

basioccipital

lamina orbitonasalis

parasphenoid

4 mm

FIGURE 3. Dorsal and ventral view of cranium. Protosalanx chinensis, CAS-SU 6306, 158 mm.

(his Salanx chinensis and Salangichthys micro-
don). He particularly noted the well-developed
fourth hypobranchials, "which so far as known
are absent from all other adult teleostean fishes."
My own observations and drawings of salangid

gill arches agree closely with Nelson's. Rosen
(1974; figs. 16g, 26a & b) figured and com-
mented briefly upon the caudal skeleton and por-
tions of the gill arches of Neosalanx brevirostris
(his Salangichthys microdori).

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

frontal

supraethmoid

parasphenoid

1 mm

FIGURE 4. Dorsal and ventral view of cranium. Salanx cuvieri, CAS-SU 32454, 69.4 mm.

The advent of a technique for staining whole
specimens with alcian and alizarin (Dingerkus
and Uhler 1977) made the present relatively ex-
tensive observations possible but even so there
have been difficulties. Some specimens stained
well with alcian but not with alizarin, or vice
versa, and in some specimens that otherwise
stained well with both stains there are still por-
tions of the skeleton which failed to take up no-
ticeable amounts of either stain. Such difficulties

could not always be made up for by staining
additional specimens.

In general, alizarin stains only bone. Alcian
stains cartilage but also stains some skeletal fea-
tures which are obviously bony and have no car-
tilaginous precursors, such as fin-rays. Cartilag-
inous structures, however, often stain much more
deeply with alcian than such non-cartilaginous
structures. Thus the salangoid hyopalatine is al-
most always stained deep blue and the opercle

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

189

ethmoid plate

supraethmoid

frontal

2 mm

parietal

FIGURE 5. Dorsal view of cranium and membrane bones on dorsal surface of cranium. Salanx prognathus, CAS 51439, 1 10
mm.

appears variably pale blue and/or red. In a few
of my figures such differences are indicated by
the intensity of stippling, but in general the dis-
tribution of stain is far too complex to permit its
representation in black-and-white illustrations.
Some idea of the difficulty involved may be gained
from Figure 20 (pelvic girdle of Protosalanx), in
which the distribution of stain is indicated. In
the cranium the distribution is far more com-
plicated and could be conveyed only by illustra-
tions in full color.

CRANIUM

(Figures 3-8)

The cranium of all salangoids is depressed,
very strongly in Salanginae and almost as strong-
ly in Protosalanginae, but relatively moderately
in Salangichthyinae and Sundasalangidae. Some
other features correlated with the cranial depres-
sion are the peculiarly underslung maxilla, ven-
trolateral eye position (especially in Salanginae),
and perhaps the posteriorly recurved jaw teeth
(especially in Salanginae and Protosalanginae).

190

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

tectum synoticum auditory capsule

cranial fontanel

foramen
' magnum

supraethmoid

taenia marginalis

epiphyseal bar semicircular canals

pila prooptica

nasal recess

basioccipital

lamina orbitonasalis

trabecula communis

hyomandibular fossa

parasphenoid

1 mm

FIGURE 6. Dorsal and ventral views of cranium. Neosalanx jordani, CAS 52028, 38.3 mm.

The development of the cranial fontanel ex-
hibits considerable differences. The fontanel ap-
parently remains open anterior and posterior to
the epiphyseal bar throughout life in Salangich-
thyinae and Sundasalangidae, although the an-
terior portion may be greatly reduced in larger
Salangichthyinae. In Protosalanginae the ante-

rior portion closes while the posterior portion
always remains open, albeit much reduced in the
largest specimens examined. In Salanginae the
cranial fontanel is entirely closed in all specimens
in which skeletal preparations have been ex-
amined.
Young Osmeridae in which the cranium is still

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

191

auditory fenestra

taenia marginalis

lamina orbitonasalis

parachordals

trabecula communis

hypophysial fenestra

notochordal groove

1 mm

FIGURE 7. Dorsal and ventral views of cranium. Sundasalanx microps, CAS 44220, 17 mm.

cartilaginous have a median bar (taenia tecti me-
dialis) separating the anterior and posterior por-
tions of the cranial fontanel into left and right
halves. Such a feature is usually but not invari-
ably absent in salangoids. In a series often Neo-
salanx jordani (39.7-45.7 mm), nine have the

cranial fontanel entirely undivided, but one (4 1 .0
mm) has a median cartilaginous bar dividing both
the anterior and posterior portions of the fon-
tanel. The bar is slender posteriorly, but ante-
riorly it is much wider, so that the anterior por-
tion of the fontanel is represented by two widely

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

4 mm

1 mm

cranial fontanel

ethmoid plate

anterior myodome

lateral fenestra

pila prooptica
trabecula communis

lamina
orbitonasalis

pila prooptica

1 mm

FIGURE 8. Lateral view of cranium, (a) Protosalanx chinensis, CAS-SU 6306, 158 mm; (b) Salanx cuvieri, CAS-SU 32454,
69.4 mm; (c) Neosalanx jordani, CAS 52028, 38.3 mm; (d) Sundasalanx microps, CAS 44220, 17 mm.

separated and relatively small openings. The epi-
physeal bar in this specimen is also larger than
usual. The condition of the cranial fontanel in
this specimen closely resembles that observed in
osmerid chondrocrania. In Protosalanx of 85-
89 mm, the anterior portion of the cranial fon-
tanel is similarly divided into greatly reduced left
and right openings, which become entirely closed
in specimens slightly larger.

The ethmoid plate is greatly enlarged in all
salangoids. In Salangichthyinae and Sundasa-
langidae it is broad and moderately elongate,
while in Protosalanginae and Salanginae it is
broad and extremely elongate. Ossification of the
chondrocranium is relatively poor in all salan-
goids but varies greatly. The greatest amount of
cranial ossification is observed in the skulls of
the largest Protosalanx, in which the supraeth-
moid, frontals, parietals, parasphenoid, and basi-
occipital are all stained more or less deeply with
alizarin. In large Protosalanx the posterior por-
tion of the parasphenoid has broad lateral wings
and the basioccipital has small thin lateral wings
(largely obscured by the overlying parasphe-
noid). Neither of these features has been ob-
served in other salangoids. In all other salangoids
the basioccipitai ossification is apparently re-
stricted to the basioccipital centrum.

In Protosalanginae the outline of the cranium
is more irregular, suggesting a more primitive
condition; while in Salanginae it is relatively
smooth and streamlined, suggesting a more de-
rived or specialized condition. The auditory cap-
sules are most pronounced or laterally prominent
in Salangichthyinae.

The interorbital septum is relatively open in
Sundasalangidae and Salangichthyinae, almost
as open in Protosalanginae, but greatly reduced
in Salanginae. In Salangichthyinae the anterior-
medial portion of the orbit is occupied by very
large pilae proopticae arising from the ventral
surface of the taenia marginalis or anterior su-
praorbital cartilage. In Sundasalangidae the pilae
proopticae are rudimentary.

A number of cranial features that occur in Sun-
dasalangidae have not been observed in the other
(mostly juvenile and adult) salangoids examined.
Thus the lamina orbitonasalis, which appears as
a single apparently simple entity in other sa-
langoids, has two components in Sundasalanx:
a dorsoanterior contribution from the taenia
marginalis and a ventroposterior contribution
from the trabecular communis or posteroventral
portion of the ethmoid plate. The ethmoid plate
is separated by the anterior myodome into dorsal
and ventral portions; the anterior myodome ex-
tends anteriorly almost to the tip of the snout.
In other salangoids the anterior myodome lies
much farther posterior, and the ethmoid plate is
relatively thin and more or less greatly depressed
(least so in Salangichthyinae).

In Sundasalanx the base of the cranium is
largely occupied by the hypophysial fenestra, a
character of all developing teleost chondrocrania
usually lost at an early stage. In all other salan-
goids the hypophysial fenestra is closed off by
cartilaginous growth and the area it once occu-
pied may be overlaid by the parasphenoid. In
Sundasalangidae the passage for the internal ca-
rotid artery is represented by an anterolateral

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

193

hyopalatine

opercle

premaxilla

supramaxilla maxj||a/

preopercle

hyomandibular

dentary

Meckel's cartilage

coronoid process
4 mm

articular

quadrate

interopercle

subopercle

FIGURE 9. Lateral view of jaws, suspensorium, and opercular bones. Protosalanx chinensis, CAS-SU 6306, 158 mm.

extension of the hypophysial fenestra; in all other
salangoids the passage for this pair of arteries is
isolated and widely separated.

In Sundasalanx the base of the cranium ex-
hibits a median groove on either side of which
extends a slight ridge. This groove probably rep-
resents the pathway of the embryonic cranial no-
tochord before its absorption (complete in all
other salangoids examined) into the basioccipital
centrum. The ridges on each side may be rem-
nants of the parachordal cartilages.
JAWS

(Figures 9- 13)

The jaws of salangoids are relatively general-
ized, in that the jaw bones, their shape, and the
distribution of teeth on them are similar to those
in many lower teleosts. In all salangoids the max-
illa is toothed and enters broadly into the gape.
All salangoids have a single supramaxilla, except
Salanginae, in which this element is lacking. In
some Salanginae the bony tip of the lower jaw is
formed not by the dentaries, but by a median
presymphyseal bone (usually tooth-bearing). Due
in part to poor quality of alcian-alizarin staining
of the lower jaw in salangoids, the relationships

of bones that constitute it have not been ade-
quately observed. The premaxillae and maxillae
are somewhat variable (see remarks in system-
atic account).

SUSPENSORIUM
(Figures 9- 13)

The outstanding feature of the salangoid sus-
pensorium is the union of the hyomandibula (hy-
osymplectic) and pterygoquadrate, which are
united into a single continuous cartilaginous ele-
ment, here called the hyopalatine (=palatohyo-
mandibuloquadrate of Roberts 1981). Only in
Sundasalanx praecox is the hyopalatine divided
into anterior and posterior portions, but the di-
vision apparently is more anterior than the prim-
itive division between hyomandibula (or hy-
osymplectic) and pterygoquadrate.

In developing vertebrates the rudimentary
mandibular arch divides into two cartilages where
it bends around the corner of the mouth: the
pterygoquadrate bar (dorsal) and the mandibular
bar or Meckel's cartilage (ventral). The rudi-
mentary hyoid arch divides into the hyoman-
dibular (dorsal) and hyoid bar (ventral). All sa-
langoids except Sundasalanx praecox show the

194

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

supraorbital

hyopalatine

pseudobranch

. 1 mm ,

FIGURE 10. Lateral and medial views of jaws, suspensorium, and opercular bones. Salanx cuvieri, CAS-SU 32454, 69.4
mm.

most unusual condition of having the dorsal por-
tions of the mandibular and hyoid arches fused
into a single element. This salangoid element has
readily definable features corresponding to the
palatine or pterygoid, quadrate, and hyoman-
dibula of teleosts in which these elements are
separate, but it is unclear whether a portion rep-
resenting the symplectic is present.

No separate symplectic has been detected in
any salangoid; the symplectic may be represented
by a thickening or ridge near the ventral margin
of the quadrate portion of the hyopalatine.

In Sundasalangidae and some Salanginae and
Salangichthyinae the suspensorium consists sole-
ly of the cartilaginous hyopalatine, but in other
Salanginae and Salangichthyinae and in Proto-
salanginae a number of perichondral, endochon-
dral, or dermal ossifications develop on the sus-
pensorium. The elements most often added are
the mesopterygoid and an anterior palatal tooth-
plate (=ectopterygoid?), which may or may not
bear teeth. The suspensorium exhibits more os-
sification in large Protosalanx than in any other
salangoids examined: heavily toothed palatal
toothplate, mesopterygoid, and partial ossifica-
tion of quadrate and hyomandibula.

Whether the dorsal portions of the mandibular
and hyoid arches are similarly fused in any other
fishes is unknown. In the few fishes for which the

development of these arches has been adequately
observed it would appear they are separate, in-
cluding Salmo (DeBeer 1937), Flops (pers. obs.),
Hepsetus (Bertmar 1959). In young salmoni-
forms I examined (including Salmo, Galaxias,
Lepidogalaxias, Hypomesus, and Spirinchus)
cartilaginous pterygoquadrate and hyomandib-
ular or hyosymplectic are always separate.

Circumorbital Bones

(Figure 12)

A supraorbital bone is seen in all Salangidae
but is absent in Sundasalangidae. The dermo-
sphenotic or sixth infraorbital appears to be ab-
sent in all salangoids. An isolated infraorbital
(fourth or fifth?) is seen in some Salangichthyinae
but is greatly reduced (Fig. 1 2).

Gill Arches

(Figures 14-17)

The upper elements of the gill arches of sa-
langoids are relatively generalized and, except in
Sundasalangidae, so are the lower elements. Ex-
cept for the upper and lower pharyngeal tooth-
plates the salangoid gill arches apparently are
entirely cartilaginous. Four basibranchials are
probably present in all salangoids but in none
are all of them separate. In Protosalanginae, Sa-

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

195

, 1 mm |
FIGURE 1 1. Lateral view of jaws, suspensorium, and opercular bones. Salanx prognathus, CAS-SU 51439, 1 10 mm.

langinae, and Salangichthyinae basibranchials 2
and 3 are indistinguishably fused to each other,
and in some Salanginae basibranchials 1 , 2, and
3 may be so fused.

All salangoids have four hypobranchials; hy-
pobranchial 4 is always separate and relatively
large.

The basibranchial series in salangoids is en-
tirely cartilaginous. Not only do the basibran-
chials themselves not ossify, but basibranchial
toothplates, a characteristic feature of many sal-
moniforms including salmonids, osmerids, and
galaxiids, are absent. The basic basibranchial ar-
rangement in Salangidae appears to be basi-
branchial 1 separate, basibranchials 2 and 3 fused,
and basibranchial 4 separate. A basibranchial 5
is fused to basibranchial 4 in various salmoni-
forms, and is apparently usually present in many
salmonoids, osmeroids, and galaxioids (includ-
ing Lepidogalaxias) as a thin cartilaginous shaft
projecting posteriorly between the fifth cerato-
branchials. In some instances there is a clear de-
marcation between basibranchials 4 and 5, and
they may be separate or at least not completely

fused. Basibranchial 5, fused with basibranchial
4, is indicated in Salangidae by Nelson (1970),
but in Salangidae I have examined there is no
indication of a fusion or demarcation between
the presumed basibranchial 5 and basibranchial
4. Basibranchial 5 does not project so far pos-
teriorly nor is it slender and rodlike as in other
Salmoniformes in which its presence is less
doubtful. I therefore tentatively consider basi-
branchial 5 absent in Salangidae. That it is absent
in Sundasalangidae seems highly likely.

Gill rakers are poorly ossifed (never stained
with alizarin) and edentulous (frequently dentig-
erous in salmonoids, osmeroids, esocoids). Those
on the trailing (inner) face of the arches usually
are fewer and smaller than those on the leading
(outer) face (Figs. 14-17). Total number of gill
rakers on leading face of first gill arch is 8-19 in
Salangidae and 0-10 in Sundasalangidae (Table
2).

Dentition

The most complete and presumably most
primitive dentition in salangoids is observed in

1 mm

FIGURE 12. Lateral view of jaws, suspensorium, and opercular bones. Neosalanx jordani, CAS 52058, 35.1 mm.

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

PL HQ

FIGURE 13. Lateral view of jaws, suspensorium, and opercular bones, (a) Sundasalanx praecox, CAS 52031, 17 mm; (b)
Sundasalanx microps, CAS 44290, 17 mm. HQ = hyomandibula + quadrate, LJ = lower jaw or Meckel's cartilage, MX =
maxilla, P = premaxilla, PHQ = hyopalatine cartilage, PL = palatine, OP = opercle, SO = subopercle.

hypobranchials

urohyal
branchiostegal rays

basibranchials

ceratobranchials

1 mm

FIGURE 14. Dorsal and ventral views of hyoid and branchial arches. Protosalanx chinensis, CAS-SU 6306, 153 mm.

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

-basihyal

dorsohyal
anterohyal

posterohyal

interhyal

basihyal

toothplate

accessory
cartilage

uncinate
process

epibranchials

197

infrapharyngobranchials

upper pharyngeal
toothplate

lower pharyngeal
toothplate

4 mm

2mm

FIGURE 1 5. Dorsal view of hyoid and branchial arches and ventral view of upper pharyngeal elements. Salanx cuvieri, CAS-
SU 32454, 69.4 mm.

, 1 mm |

FIGURE 16. Dorsal view of hyoid and branchial arches; ventral view of infrapharyngobranchial 4 and upper pharyngeal
toothplate. Neosalanx jordani, CAS 52058, 38.3 mm.

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

basibranchial 1 + 2 +
hypobranchial 1 + 2

basibranchial 3 +
hypobranchial 3

hypobranchial 4
basibranchial 4

, 1 mm

FIGURE 17. Dorsal view of hyoid and branchial arches. Sundasalanx microps, CAS 44220, 17 mm.

Protosalanx, with numerous large, conical teeth
on the premaxilla, maxilla, palatal toothplate
(=ectopterygoid?), tongue (basihyal toothplate),
and upper and lower pharyngeal toothplates. As
in all Salangidae, there are only two pairs of pha-
ryngeal toothplates: the upper, on infrapharyn-
gobranchial 4, and the lower, on ceratobranchial
5; the branchial arches are otherwise entirely
toothless.

In Protosalanx the teeth on the palatal tooth-
plate and lower jaw are in two rows, as in many
other salmoniforms, but in all other salangoids
the teeth on each tooth-bearing element except
those in the pharynx are restricted to single rows.
In largest specimens of Protosalanx the tongue
teeth are more or less widely scattered over the
surface of the basihyal (as in Fig. 9), but in small-
er ones they are restricted to two marginal rows,
as in salmoniforms generally. The only other sa-
langoid with tongue teeth, Salanx (Leucosoma)
reevesi, has them in a single median row on the
basihyal toothplate, a unique specialization for
salmoniforms. This character is diagnostic of the
subgenus Leucosoma.

The maxilla and lower jaw are well-toothed in
all salangoids; the palate is toothless in Neosa-
lanx and Sundasalanx. In Neosalanx the teeth
on the premaxilla, maxilla, and lower jaw are
very small, and frequently the premaxilla and
lower jaw are entirely toothless. In Sundasalanx
bony pharyngeal toothplates apparently fail to

form, and the pharyngeal teeth appear to be di-
rectly attached to the cartilaginous infrapha-
ryngobranchial 4 and ceratobranchial 5. The only
bony tooth-bearing elements in Sundasalanx ap-
pear to be the premaxilla and maxilla; the lower
jaw teeth are loosely attached to Meckel's car-
tilage.

PECTORAL GIRDLE

(Figures 18-19)

All salangoids have a secondary pectoral girdle
(connecting the primary girdle to the back of the
cranium) consisting of three dermal bones: post-
temporal, supracleithrum, and cleithrum. Post-
cleithra are absent except in Salanginae, in which
there is a single postcleithrum. In Salangidae, the
primary shoulder girdle consists of the entirely
cartilaginous paired scapulocoracoids and one or
two series of radials. The basic number of pri-
mary radials appears to be five in all Salangidae.
The first primary radial, associated with the out-
ermost (enlarged) pectoral fin-ray, is relatively
simple; it is largest in males of Protosalanginae
and Salanginae. The other primary radials are
complex, with numerous deep divisions approx-
imately corresponding in number to the fin-rays.
These divisions are most numerous in Salangich-
thyinae, particularly Neosalanx, but are well de-
veloped in all Salangidae. Comparable divisions
or fimbriae occur in the pectoral basal plate of

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

199

post temporal

supracleithrum

proximal radials1-5

distal radials

postcleithrum

1 mm

FIGURE 18. Left half of pectoral girdle, (a) Protosalanx chinensis, CAS-SU 6306, 158 mm (medial view); (b) Salanx cuvieri,
CAS-SU 32454, 69.4 mm (lateral view); (c) Neosalanxjordani, CAS 52058, 43. 1 mm (dorsal view); (d) Salangichthys ishikawae,
CAS 6780, 74 mm (lateral view).

the salmoniform Dallia pectoralis but are not
present in other salmoniforms I have examined
and do not seem to have been reported in any
other teleosts. Secondary radials, more or less
corresponding in number to the pectoral fin-rays,
are small and simple. The mesocoracoid is lack-
ing in all salangoids except that Protosalanx has
a process on the median surface of the scapu-
locoracoid that may represent the ventral portion
of the mesocoracoid (Fig. 1 8a, medial process).
In Sundasalangidae the primary pectoral girdle
consists of a U-shaped median scapulocoracoid
and a basal plate. Fin-rays are absent.

PELVIC GIRDLE

(Figure 20)

The left and right halves of the pelvic girdle
develop in the ventral myotomic wall, and, as
the ventral myotomic progression is arrested in
Salangidae while the myotomes are still widely
separated, the pelvic girdle halves remain widely
apart and fail to form any sort of ligamentous or
cartilaginous connection between each other. As
pointed out by Klyukanov (1975), in Salmoni-

formes the two halves of the pelvic girdle are
usually joined at least anteriorly for a short dis-
tance by strong cartilaginous or ligamentous tis-
sues.

AXIAL SKELETON

(Figure 1)

All salangoids have a pair of small dorsal car-
tilages straddling the intervertebral disc between
the basiocciptal and first vertebral disc; such car-
tilages occur in many (perhaps most or all) Sal-
moniformes.

In all Salangidae the neural arches of vertebrae
1 and 2 are fused dorsally; this condition has not
been observed in Osmeridae or any other sal-
moniforms I have examined. In Sundasalangidae
the neural arches of vertebrae 1 and 2 are sep-
arate from each other and morphologically sim-
ilar to those of the vertebrae immediately suc-
ceeding them.

In salangoids the mineralized portion of each
centrum is relatively elongated and hourglass
shaped, so that the intervertebral joints are nar-
row and the notochord greatly constricted. In

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

1 mm

FIGURE 19. Pectoral girdle, (a) Sundasalanx microps, CAS 44220, 17-mm adult (posterior view); (b) Elops hawaiiensis, CAS
52035, 30-mm leptocephalus larva (posterior view); (c) Sardina pilchardus, 20-30 mm (ventral view?, after Goodrich 1922);
(d) Dallia pectoralis, (lateral view, after Starks 1904; apparently based on CAS-SU 12615, 125 mm, Nushagak River, Alaska);
AP = ascending process, CL = cleithrum, F = fin margin, PP = posterior process, PT = posttemporal, R = basal plate, SCL =
supracleithrum, SCO = scapulocoracoid. In (b) and (c) the first primary radial has pinched off from the basal plate.

salmonids, osmerids, galaxiids, and other Sal-
moniformes, especially in the young stages, the
mineralized portion of each centrum tends to be
relatively short and cylindrical, so that the in-
tervertebral space is much larger and the verte-
bral section of the notochord is entirely intact.
A comparable condition is not present in any
salangoid skeletal material I have examined.

Ribs are absent or weakly developed and stain
poorly. They are small, weakly stained with al-
cian when present (Fig. Ib).

Gosline (1960) and others have pointed out
that neural and hemal spines of most Salmoni-

formes, especially posteriorly, may be flattened
or laminar, even to the extent of resembling a
continuous keel. The neural and hemal spines of
salangoids are always relatively slender, espe-
cially posteriorly.

A round, oval, or elongate and splintlike adi-
pose fin cartilage lies at the base of the adipose
fin in all Salangidae. A survey of lower teleosts
for the adipose fin cartilage by Matsuoka and
Iwai (1983) revealed its presence in Salangidae,
Osmeridae, Plecoglossidae, Myctophidae, and
Neoscopelidae; it was not observed in other low-
er teleosts with an adipose fin including Sal-

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

201

radials1-4

blue

1 mm

blue

parapelvic
cartilages

1 mm

FIGURE 20. Ventral view of left half of pelvic girdle, (a) Protosalanx chinensis, CAS-SU 6306, 158 mm; (b) Salanx cuvieri,
CAS-SU 32454, 69.4 mm; (c) Neosalanx jordani, CAS 52028, 43.1 mm; (d) Sundasalanx microps, CAS 44220, 17 mm (with
lateral view of pelvic girdle and parapelvic cartilages above).

monidae, Retropinnidae, Prototroctidae, Aulo-
podidae, Synodontidae, Chlorophthalmidae,
Argentinoidei, Characoidei, or Siluriformes. The
similar morphology of the adipose fin cartilages
in Salangidae and Osmeridae, as noted by Mat-
suoka and Iwai, is possibly indicative of rela-
tionship between these two families.

Caudal Fin Skeleton

(Figure 21)

The caudal fin is more or less deeply forked,
and the upper and lower lobes are about equal.
Principal caudal fin-rays are invariably 10 + 9;
upper and lower procurrent caudal fin-rays are
moderately numerous (to 14). The complex ural
or hypural centrum apparently consists of three
centra and uroneural 1 (sometimes also uro-
neural 2?) fused into a single unit. The three cen-
tra involved are the terminal centrum and post-
terminal centra 1-2, according to the
nomenclature of Gosline (1960), or preural cen-
trum 1 and ural centra 1-2, according to Rosen
( 1 974). In none of the skeletal material examined
is there any indication of separate centra poste-
rior to the complex hypural centrum. Epurals 0-

3. A separate uroneural 2 is sometimes present,
but uroneural 1 is apparently always fused to
complex hypural centrum. Free radial or ptery-
gial cartilages are sometimes present, usually be-
tween ray halves at the base of the anteriormost
2-3 upper or lower procurrent rays and the low-
ermost upper and uppermost lower principal rays.
Hypurals six. Six separate hypurals occur in Sa-
langichthys microdon (Rosen 1 974, Fig. 26). Pro-
tosalanx chinensis occurs with hypurals 1-2 and
5-6 separate, but with 3-4 fused near the base.
The hypurals are more fused in Neosalanx, Sa-
lanx, and Sundasalanx. In Salanx parhypural
and hypurals 1-2 are fused near the base; hy-
purals 1-2 and 3-4 are fused for their entire length
except for oblong basal foramina where fusion
evidently failed to complete. In Sundasalanx
parhypural and hypurals 1-3 are evidently fused
into a single element.

SYSTEMATICS

In the present account the salangoids are rec-
ognized as a salmoniform superfamily separate
from osmeroids, which they superficially resem-
ble. There are two families, Sundasalangidae, with

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

uroneural epura|
ural centrum

opisthural

hypurals1-6

neural spines

hemal spines
hypurapophysis

accessory
cartilages

1 mm

parhypural

1 mm

FIGURE 21. Lateral view of caudal fin skeleton, (a) Protosalanx chinensis, CAS-SU 6306, 158 mm; (b) Salanx cuvieri, CAS-
SU 32454, 61.7 mm; (c) Neosalanx jordani, CAS 52028, 43.1 mm (note: hypurals 2 and 3, normally separate from each other
in all salangoids, are fused in this specimen); (d) Sundasalanx microps CAS 44220, 17 mm. In a-c left half of fin rays removed
to facilitate observation of median structures.

only a single genus and two species, and Salangi-
dae. Salangidae is further divided into three
subfamilies, four genera, and eleven species. The
genus Salanx is further divided into three sub-
genera; this taxonomic category is not employed
in the other genera of salangoids. In addition to
the new superfamily Salangoidea, the new
subfamily Salangichthyinae is proposed for Neo-
salanx and Salangichthys, leaving the subfamily
Protosalanginae with only the genus Protosa-
lanx. No new genera or species are proposed.
Some previous workers, particularly Regan
(1908b) and Fang (1934a, b) recognized far more
species than I have, especially in the subgenus
Salanx (genera Salanx and Parasalanx of Re-
gan). This is attributable in part to their basing
species on only one or a few type-specimens and
utilizing characters such as cranial proportions,
body depth, and relative position of dorsal and

anal fins which vary considerably within the
species. Neither Regan nor Fang utilized verte-
bral counts, which I find extremely useful in dis-
tinguishing species. My extensive data on ver-
tebral counts of types and other material are
presented in Table 2.

My counts of vertebrae, fin-rays (except pelvic)
anal scales, branchiostegal rays, and gill rakers
are presented in Table 2. This table includes all
species of salangoids herein recognized as valid
except Neosalanx reganius, which I have not ex-
amined. Pelvic fin-ray counts are excluded be-
cause they are invariably 5 in Sundasalangidae
and almost invariably 7 in Salangidae (6 in one
observed specimen of Neosalanx jordani, 8 in
two specimens of Salangichthys microdori). Pre-
vious authors have presented data on most of
the species but have often lumped data from
various localities (and frequently of two or more

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

203

species). In order to minimize this problem, my
data are presented separately for each locality.

Although Table 2 includes meristic data ob-
tained from whole specimens, radiographs, and
cleared and stained specimens, the stained spec-
imens provide the best material for accurate count
of fin-rays, teeth, and gill rakers. It is difficult to
observe the jaw teeth and lowermost pectoral fin-
rays in Neosalanx except in stained material. In
dorsal and anal fin-ray counts the last "two" rays
("divided to base") are counted as one ray. In
vertebral counts the basioccipital centrum is not
counted and the hypural complex centrum is
counted as one. In salangoids, especially in fe-
males, the anteriormost anal-fin pterygiophore
does not provide a ready basis for distinguishing
abdominal and caudal vertebrae. In order to ob-
tain additional meristic data from the vertebral
column and at the same time obtain more precise
data on the relative position of fin origin, I have
taken data on the vertebrae parallel to the origins
of the pelvic, dorsal, and anal fins. The number
of vertebrae posterior to a vertical line through
the base of the last anal fin-ray is also recorded.

Radiographs are usually satisfactory for ob-
taining vertebral counts of salangoids and some-
times for fin-ray counts. Sometimes the verte-
brae may show up very faintly but it is almost
always possible to obtain a count repeatable to
within one vertebra. Fin-rays, however, fre-
quently cannot be accurately counted on radio-
graphs, and I have only incorporated data on
fin-ray counts taken from radiographs when the
radiographs seemed reliable.

Some characters utilized by other workers to
distinguish species are not emphasized here be-
cause they do not seem useful. This particularly
applies to pectoral fin-ray counts in Neosalanx
and to the elongation of the head or cranium,
relative position of the dorsal- and anal-fin bases,
and body depth, especially in Salanx. In salan-
gids the number of pectoral fin-rays generally
continues to increase slightly with growth, es-
pecially so in those such as Neosalanx, in which
the rays are exceptionally numerous. The elon-
gation of the cranium (particularly its anterior
portion) is extremely variable in Salanx, as not-
ed also by Wakiya and Takahasi (1937:289). This
variation is individual and is probably enhanced
by growth. The position of the dorsal and anal
fins relative to each other is also highly variable
in salangids, subject to individual variation as

well as sexual dimorphism. In defining species
of Salanx too much reliance has been placed on
slight differences in fin positions based on only
one or two specimens. Salanx, Salangichthys,
and other salangids vary enormously in body
depth due to sex-related body changes and non-
sexual factors of condition and preservation.

In discussing salmonoid classification, Gosline
(197 1:1 19) stated:

The suborder Salmonoidei as here recognized (Families Sal-
monidae, Osmeridae, Plecoglossidae, Salangidae, Retropin-
nidae, Aplochitonidae, and Galaxiidae) is a group of highly
diverse inshore and freshwater salmoniform fishes. Though
the included families no doubt should be divided into su-
perfamily groupings, inadequate knowledge of the Salan-
gidae and the Southern Hemisphere forms would seem to
make any formal superfamily classification premature at
the present time. Informally, the members may be divided
between Northern and Southern Hemisphere forms. The
diverse forms from the Southern Hemisphere seem to be
most closely related to the northern osmerids. . . . The
Northern osmeroids are represented by four quite distinct
lines: Salangidae, Plecoglossidae, Osmeridae, and Salmon-
idae.

Rosen (1974) divided the suborder Salmo-
noidei into two superfamilies, Salmonoidea— in-
cluding the Southern Hemisphere families (ex-
cept Retropinnidae) and Salmonidae — and
Osmeroidea (with four families listed as incertae
sedis: Osmeridae, Plecoglossidae, Retropinni-
dae, and Salangidae). I have not investigated
Retropinnidae or the highly aberrant Plecoglos-
sidae but suspect that Retropinnidae (particu-
larly Prototroctes) and Plecoglossus may indeed
be closely related to each other and perhaps to
Osmeridae. But I have not been able to find any
good evidence (in the form of shared specializa-
tions or derived characters) between Salangidae
and any one or combination of these families. I
have therefore designated the new superfamily
Salangoidea, which is coequal with the superfam-
ilies Osmeroidea and Salmonoidea (and Galax-
ioidea, if this is also to be recognized).

SALANGOIDEA, NEW SUPERFAMILY

This superfamily apparently differs from all
other Pisces in having a suspensorium in which
the cartilaginous palatine and pterygoid (of the
mandibular arch) and quadrate and hyomandib-
ular (of the hyomandibular arch) are fused into
a single element, the hyopalatine. Gill arches with
well-developed fourth hypobranchials— so far as
known absent from all other adult teleosts (Nel-

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

son 1970). Pharyngobranchials 4. Anterior pha-
ryngobranchial modified as an elongate "suspen-
sory pharyngeal"; only fourth pharyngobranchial
bears teeth (teeth absent in Sundasalanx). Fifth
ceratobranchial with well-developed teeth (ab-
sent in osmeroids; McAllister 1963:4). Bran-
chiostegal rays 2-5 (7-19 in salmonoids, 5-10
in osmeroids, 3-9 in galaxioids). Cranium mod-
erately to excessively flattened (more so than in
any other salmoniforms). Maxillary bone, bear-
ing teeth for its entire length, with its posterior
half abruptly curved medially beneath head (so
that teeth on posterior half of maxillary are di-
rected anteriorly rather than ventrally). Scales
entirely absent except for a row of strongly ad-
herent anal scales in adult male Salangidae.

Dermosphenotic and circumorbital bones ab-
sent, except for a single small troughlike bony
element observed in Neosalanx, which may rep-
resent a fifth or sixth circumorbital (not dermo-
sphenotic). Supraocciptal bone absent (present
in most other salmoniforms).

Pectoral fins pedunculate throughout life (with
pectoral radials in a fleshy pedestal separate from
body). Pelvic fin-rays usually 5 or 7 (rarely 6 or
8; 8 in osmeroids). Principal caudal fin-rays in-
variably 10 + 9 (as in most lower teleosts includ-
ing salmoniforms with generalized caudal fins;
galaxioids have fewer).

Salangoids apparently have no laterosensory
canals on the body. The cephalic laterosensory
canals, although well developed, are superficial
(i.e., not enclosed in bony tubules) and often dif-
ficult to observe in their entirety. Those of Sa-
lanx chinensis, illustrated by Nelson (1970, Fig.
1 5), do not exhibit any particularly unusual fea-
tures for lower teleosts. There are preopercular,
mandibular, supraorbital, infraorbital, and ex-
trascapular canals. The mandibular is not con-
tinuous with the preopercular. The supraorbital
and infraorbital extend anteriorly only a short
distance in front of the nostrils, i.e., not signifi-
cantly onto the greatly depressed and enlarged
snout. The infraorbital has 8 pores, the preoper-
cular 6, and the mandibular 5.

Alimentary canal a relatively simple, straight
tube. Pyloric caecae absent. Gonads paired.

Salangidae Jordan and Snyder, 1 902

Pelvic fin almost invariably with 7 rays (8 ob-
served in one specimen of Salangichthys ishi-
kawae and two S. microdon, 6 in one Neosalanx

jordani). Pelvic girdle without parapelvic carti-
lages. Pectoral fin-rays 8-34. Pectoral girdle with
five proximal radials; distal ends of one or more
proximal radials with more or less numerous
branches; adult males with a series of anal scales
and enlarged, modified anal fins; total vertebrae
49-79.

The family Salangidae comprises three
subfamilies: Protosalanginae, Salangichthyinae,
and Salanginae.

Protosalanginae Wakiya and Takahasi, 1937

This subfamily, here restricted to the mono-
typic genus Protosalanx, differs from all other
salangoids in having the premaxilla, palatal
toothplate (=ectopterygoid?), and dentary with
two rows of teeth instead of at most a single row;
the basihyal toothplate of the tongue also has the
teeth in two marginal rows (a primitive condition
for salmoniforms) or irregularly scattered over
its entire surface; the only other salangoid with
basihyal teeth has them in a single median row.
Pelvic fins relatively larger and more anterior
than in any other salangoids (see Fig. 1, Table

2).

Cranium strongly depressed (almost as much
as in Salanginae); adults with anterior portion of
cranial fontanel closed, posterior portion of cra-
nial fontanel greatly reduced but remaining open
throughout life (both portions closed in adult
Salanginae, open throughout life in Salangich-
thyinae and Sundasalangidae). Lower jaw weakly
projecting beyond upper jaw; premaxillae pro-
jecting anteriorly beyond snout tip as in Sa-
langinae but failing to form a membrane-covered
space through which symphyseal teeth of lower
jaw project. Lower jaw without enlarged sym-
physeal teeth (present in Salanginae), sometimes
with a weakly developed fleshy presymphyseal
process but without presymphyseal teeth or bony
process. Adults attaining slightly greater stan-
dard length (Table 1) and heavier-bodied than
any other salangoids. Dorsal fin-rays 16-18 and
anal fin-rays 30-32 (vs. 10-15 and 14-32 in all
other salangoids); vertebrae 66-70 (Table 2).

Protosalanx Regan, 1 908

Eperlanus BASILEWSKY, 1855:242.

Salanx ABBOTT, 1901:490.

Protosalanx REGAN, 1908b:444 (type-species, by monotypy,

Salanx hyalocranius ABBOTT, 1 90 1 = Eperlanus chinensis

BASILEWSKY, 1855).
Paraprotosalanx FANG, 1934a:246 (type-species, by mono-

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

205

typy, Protosalanx andersoni FANG, 1934a (non RENDAHL,
1923) = Protosalanx chinensis BASILEWSKY, 1855).

Protosalanx chinensis (Basilewsky, 1855)

Eperlanus chinensis BASILEWSKY, 1855:242 (type-locality "in
sinu Tschiliensis habitat" [not "Pekin" as usually cited]).

Salanx hyalocranius ABBOTT, 1901:3490 (type-locality Pei-ho
at Tien-tsin).

Protosalanx hyalocranius REGAN, 1908b:445.

Paraprotosalanx andersoni FANG, 1934a:246 (Figs. 4-6, text
in part [non Paraprotosalanx andersoni RENDAHL, 1923]).

Protosalanx chinensis CHYUNG, 1961:163.

MATERIAL EXAMINED. -BMNH 1929.2.5.2.-3, 61-65 mm,
Kiangyin; CAS 52026, 60:48.2-70.7 mm, no locality (pur-
chased in San Francisco); CAS-SU 6306, 25:80.5-163 mm,
Pei-ho at Tien-tsin, paratypes of Salanx hyalocranius (7:85.5-
158 mm alcian-alizarin); CAS-SU 23639, 1:137 mm, Seoul;
CAS-SU 36025, 3:120-136 mm, no locality; UMMZ 180096,
2:127-129 mm, Korea; USNM 120746, 2:129-132 mm, Ko-
rea.

Protosalanx appears to be the most primitive
salangoid. There is no indication that it com-
prises more than a single species. Although Ab-
bott's account begins "Salanx hyalocranius new
species," it concludes "this species is probably
identical with Eperlanus chinensis Basilewsky,
from Pekin, but the name chinensis is already
used for the 'whitebait of Makao' " (Abbott 1 90 1 :
490-491). In Abbott's time Salangidae were so
poorly known it was reasonable for him to as-
sume that his material might represent an un-
described species, but even so it is clear from this
statement that Abbott was really proposing a re-
placement name. Now that Salangidae are better
known it seems Basilewsky's account could only
refer to this species, as explicitly recognized by
Wakiya and Takahasi (1937), although they re-
tained the name P. hyalocranius. The holotype
of P. chinensis cannot be found (Barsukov, pers.
comm. 1983). Since the "whitebait of Makao"
has been referred to as Leucosoma or Salanx
chinensis but never as Eperlanus or Protosalanx
chinensis, the epithet chinensis is available for a
species of Protosalanx. As this is also the earliest
name proposed it must replace hyalocranius, and
the species should be known as Protosalanx chi-
nensis. The only publication to come to my at-
tention in which this name is correctly applied
is by Chyung(1961).

Wakiya and Takahasi (1937) correctly iden-
tified Paraprotosalanx andersoni Fang, 1934a
with this species. Fang's figures agree in every
respect with P. chinensis. The fleshy presymphy-
seal appendage, presumed by Fang to differen-

tiate his Paraprotosalanx from Protosalanx, is
also present in some of Abbott's type-specimens
of S. hyalocranius. Fang's figures presumably are
based upon the single large male, "S. 4374," 153
mm (total length according to Table 4, but stan-
dard length according to p. 247) from Nanking.
All or almost all of the other specimens referred
to Paraprotosalanx andersoni by Fang are prob-
ably Neosalanx.

It should be noted that small specimens in
museum collections identified as Protosalanx are
usually Neosalanx and that all or almost all pub-
lished reports of smaller Protosalanx up to the
present time are based on Neosalanx. For ex-
ample, I find that all of the small specimens in
Abbott's type-series of S. hyalocranius are Neo-
salanx. Young P. chinensis are relatively rare in
collections. Those I examined (smallest 48.2 mm)
closely resemble the largest adults in every way
except they lack the sexually dimorphic char-
acters of adult males. The strongly pointed snout
and large teeth arranged in two rows on the pal-
ate, tongue, and lower jaw are easily observable.
Neosalanx have no teeth on the tongue or palate,
and the jaw teeth except on the maxillary are
absent or minute and difficult to observe, while
the males are sexually mature and provided with
greatly enlarged anal fins and anal scales at rel-
atively small size. The smallest male Protosalanx
with anal scales is probably considerably larger
than any Neosalanx.

Protosalanx chinensis appears heavier-bodied
at all sizes and to attain a greater size than any
other salangoid. The 163-mm specimen is the
largest that has been reported.

Salanginae Regan, 1908b

Cranium and especially ethmoid plate very
strongly depressed and elongate, more so than in
any other salmoniforms. Adults with cranial fon-
tanel entirely closed (posterior and sometimes
also anterior portion of cranial fontanel open
throughout life in all other salangoids). Upper
and lower jaws with strongly pointed or project-
ing tips. Teeth relatively large and few in num-
ber. Premaxillae projecting beyond concave an-
terior margin of ethmoid plate to form a
membrane-covered space penetrated by enlarged
symphyseal teeth of lower jaw. Lower jaw often
with a fleshy or bony presymphyseal process and
presymphyseal teeth (Wakiya and Takahasi
1937, pi. 20, figs. 31-34). Supramaxilla absent

206

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

(present in all other salangoids). Pectoral fin-rays
7-1 1 (20-32 in all other Salangidae). Pectoral
girdle with a single postcleithrum (absent in all
other salangoids). Body extremely elongate, more
so than in any other salmoniforms. Basal portion
of neural and hemal arches expanded, frequently
covering centrum laterally and fusing with each
other. Distal portion of neural arches with an-
terior and posterior projections, those of succes-
sive vertebrae articulating with each other. Ver-
tebrae 68-79 (37-70 in all other salangoids).

The enlarged symphyseal teeth of the dentary
and the membrane in the roof of the mouth
through which they project presumably form a
puncturing device to kill prey. Similar devices,
with foramina in the roof of the mouth through
which fanglike lower jaw teeth pass, occur in
other piscivorous or predatory teleosts, including
the characoids Hepsetus, Hop lias, and Acestro-
rhynchus (Roberts 1969).

In many respects Salanginae appear to be the
most highly specialized members of the family.
Even the low pectoral fin-ray count, which might
be considered primitive, may be secondarily
evolved, since primary pectoral radials 2-4 of
Salanginae exhibit the distally fimbriate or dig-
itate condition that is probably associated with
the retention of pedunculate pectoral fins (prob-
ably a neotenic character) and evolution of large
numbers of pectoral fin-rays (a specialization
shared by all other Salangidae).

SalanxOken, 1817

"Les Salanx" CUVIER, 1817:185 (French vernacular; not avail-
able for zoological nomenclature).

Salanx OKEN, 1817:1183 (Latinization of Cuvier's "Les Sa-
lanx," and the earliest name available for zoological no-
menclature; see ICZN Declaration 87, paragraph 12. Type-
species, by monotypy, Salanx cuvieri VALENCIENNES, 1 849).

Leucosoma GRAY, 1831:4 (type-species, by monotypy, Leu-
cosoma reevesi GRAY, 1831).

Hemisalanx REGAN, 1908b:444 (type-species, by monotypy,
Hemisalanx prognathus REGAN, 1 908b).

Parasalanx REGAN, 1908b:444 (type-species, by subsequent
designation of FANG, 1934a:259, Parasalanx gracillimus
REGAN, 1 908b = ISalanx cuvieri VALENCIENNES, 1 849).

Reganisalanx FANG, 1934b:509 (type-species, by monotypy,
Reganisalanx normani FANG, I934b = Salanx ariakensis
KJSHINOUYE, 1901).

Metasalanx WAKIYA AND TAKAHASI, 1937:293 (type-species,
by monotypy, Metasalanx coreanus WAKIYA AND TAKAHASI,
1937, a nomen nudum).

The four species herein recognized as consti-
tuting the genus Salanx have been placed by
other authors in three genera, Salanx, Hemisa-

lanx, and Leucosoma. Wakiya and Takahasi
(1937) even placed Hemisalanx in a subfamily
of its own, Hemisalanginae, regarded by them as
intermediate between Protosalanginae and Sa-
langinae. Because these four species differ strik-
ingly from all other salangids in several features
of skeletal anatomy but agree closely with each
other in conformation of the cranium and jaws,
distribution and size of jaw teeth, number of
pectoral fin-rays, and the peculiar modification
of their neural and hemal arches and high ver-
tebral counts, I prefer to recognize them as be-
longing to three subgenera in the sole genus of
the subfamily Salanginae.

Salanx (Salanx) ariakensis
(Kishinouye, 1901)

Salanx ariakensis KJSHINOUYE, 1 90 1 :359 (type-locality Ariake

Bay, Kiushiu).
Salanx acuticeps REGAN, 1908a:360 (type-locality Lake Can-

didius, Formosa).

Parasalanx acuticeps REGAN, 1908b:446.
Parasalanx longianalis REGAN, 1 908b:446 (type-locality Liao-

ho, northern China).
Parasalanx annitae VAN DAM, 1926:342 (type-locality Pei-

taiho, China).
Reganisalanx normani FANG, 1 934b:509 (type-locality Ichang,

as herein restricted).

MATERIAL EXAMINED. -AMNH 10327, 7:125-147 mm,
Hunan; BMNH 1888.5.15.1 1-12, 2:141-143 mm, Ichang (lec-
totypeandparalectotypeofT?. normani); BMNH 1898.2.8.20-
23, 4:114-123 mm, Liao-ho, northern China (syntypes of P.
longianalis); BMNH 1904.4.2835-36, 2:116-118 mm, Lake
Candidius, Formosa (syntypes of S. acuticeps); BMNH
1927.3.26.3, 125 mm, Nanking; BMNH 1928.6.22.6, 1 15 mm,
Wenchow; CAS-SU 8574, 2:99.1-104 mm, Ariake Sea (iden-
tified by Kishinouye); CAS-SU 23103, 107 mm, Maruyama,
Taihoku, Formosa; ZMA 1 12.587, 128 mm, Peitaiho, China
(holotype of P. annitae).

In vertebral counts and in all other respects so
far as known the four syntypes of P. longianalis
agree well with other material herein referred to
as Salanx ariakensis, except for their consis-
tently high anal fin-ray counts of 30-32 (reported
by Regan 1908b:446). Most samples of S. ari-
akensis examined have only 26-29 anal fin-rays,
but two specimens from Ariake Bay have 27 and
31.

Reganisalanx normani is based primarily on
the description by Regan (1908b) and supple-
mentary notes by Fang (1934b:509) of two spec-
imens from Ichang (BMNH 1888.5.15, 11-12),
identified by Regan (ibid.) as Salanx cuvieri. Fang
declared that the specimens represented a dis-
tinct genus but did not provide a proper generic

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

207

diagnosis or description; apparently he distin-
guished it from Salanx based on the lack of a
presymphyseal bone. In my opinion the char-
acter cannot be used to split the genus Salanx.

I have reidentified these specimens as S. ari-
akensis, a species in which the presymphyseal
bone may be present or absent. I have not seen
the third specimen referred to R. normani by
Fang (ibid.). It is clear from Fang's account that
he did not compare this specimen directly with
the two specimens from Ichang, and it might not
be conspecific. In order to fix the identity of this
nominal taxon, the 141 -mm undamaged speci-
men from Ichang (BMNH1888.5.15.11)is here-
by designated the lectotype. The 143-mm spec-
imen, with the body damaged just behind the
head and at mid-abdomen, is a conspecific para-
lectotype (BMNH 1888.5.15.12).

Fang (1934a) reported 1 1 specimens (as Para-
salanx longianalis) with the following anal fin-
ray counts: 28(5), 29(2), 30(3), 32(1). The ver-
tebral counts are unknown for these specimens
but it seems likely from Fang's account that they
are all S. cuvieri.

The holotype of P. annitae has the head rel-
atively short and broad (for the subgenus Salanx)
and in this respect is more like S. ariakensis than
S. cuvieri. A presymphyseal bone is present, but
it is short considering the large size of the spec-
imen, and has only 2 teeth on each side. The
premaxilla has 7 teeth, maxilla 1 2, and dentary
about 10.

Salanx (Salanx) cuvieri Valenciennes, 1 849

Salanx cuvieri VALENCIENNES in CUVIER AND VALENCIENNES,
1849:360 (type-locality unknown).

^Parasalanx gracillimus REGAN, 1 908b:446 (type-locality
Shanghai).

Parasalanx angusticeps REGAN, 1908b:446 (type-locality
China).

Parasalanx cantonensis HERRE, 1932:425 (type-locality Can-
ton).

MATERIAL EXAMINED. -AMNH 51689, 3:88.6-106 mm,
Canton; BMNH 1855.9.19.1539, 144 mm (holotype of P. an-
gusticeps); BMNH 1891.1.31 .20, 1 1 1 mm, Shanghai (holotype
of P. gracillimus); BMNH 1936.10.7.13, 1 19 mm, Sharp Peak,
Fukien; CAS 52057, 4:76.5-98.0 mm, Hong Kong (1 alizarin);
CAS-SU 225732, 112 mm, Canton (holotype of P. cantonen-
sis); CAS-SU 32454, 18:56-66 mm, Chuan Is. (4:61.7-69.4
mm alcian-alizarin); CAS-SU 32943, 117 mm, near Pakhoi,
SW Kwangtung; MNHN 9900, 1 12 mm, no locality (holotype).

So far as I have been able to determine, vari-
ation in the presymphyseal bone within each
species, including its presence or absence and its

length or amount of dentition, is correlated chief-
ly with size and is not sexually dimorphic.

NOTES ON HOLOTYPE.— The holotype (Fig. 2a)
is dried but complete and in fair condition. The
body immediately posterior to the head is badly
damaged and fin-rays brittle, so it must be han-
dled with care. Cranial width (at anterior margin
of eyes) 3.5 in cranial length. Presymphyseal bone,
2.1 mm long, with 1-2 moderately large teeth
basally and at least 2 minute teeth distally. Pre-
maxilla considerably elongated anteriorly, with
7-8 teeth. Maxilla with about 7 teeth. Dentary
with about 1 3 teeth of variable size. Palatal teeth
7, very small and in a single row. The following
proportional measurements are expressed as
times in standard length. Length of cranium about
7; length of head (to end of gill cover) 4.7; length
from anterior midline of ethmoid plate (concave)
to anterior rim of orbit 16; length from tip of
upper jaw (premaxilla) to anterior rim of orbit
1 0; diameter of eye (slightly shrunken) approx-
imately 28.

NOTES ON SYNONYMY.— P. angusticeps is dis-
tinguished by Regan primarily on the basis of its
exceptionally elongate head: "head nearly 4 times
as long as broad; snout a little longer than post-
orbital length of head" versus head 3 times or a
little more than 3 times as long as broad, and
snout only as long as or a little shorter than post-
orbital length of head in all other Parasalanx
and Salanx (Regan 1908b:445-446). The den-
tition of the holotype of P. angusticeps, a gravid
female of 144 mm, is complete and undamaged.
Presymphyseal bone elongate with 5-6 teeth on
each side; premaxilla with 7 teeth; maxillary teeth
10 or 1 1; dentary with a small tooth anteriorly
(just behind symphysis), then an enormous ca-
nine tooth, followed by 7 small teeth and 6 mod-
erately large teeth; palatine with 8 small teeth in
a single straight row.

The holotype of P. gracillimus is in poor con-
dition, dried, twisted, and slightly shrunken. Its
body depth, reported as 18 times its length, is
attributable to the poor condition (emaciation)
of the specimen. Its dentition is as follows: pre-
symphyseal bone with 3 teeth on each side, pre-
maxillary 5, maxillary 8, dentary with 1 mod-
erately large, 6 small, and 5 moderately large,
and palatal 7 moderately large. The vertebral
column is broken anteriorly, making all of the
counts based on vertebrae doubtful. Wakiya and
Takahasi (1937:288) tentatively placed P. gra-

208

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

cillimus as a synonym of P. ariakensis, and per-
haps they were correct. This matter is of some
nomenclatural significance, since P. gracillimus
is type-species of Regan's Parasalanx.

The holotype of P. cantonensis has a very elon-
gate presymphyseal bone with 10 teeth; premax-
illary teeth 8, maxillary teeth 10, dentary teeth
13; palatal teeth 11.

Length of the head (and cranium) is excep-
tionally variable in all genera of Salangidae, even
including the relatively short-snouted genus
Neosalanx, and is particularly variable in the
long-snouted subfamily Salanginae. Variable
elongation of the head and particularly the snout
with its flattened ethmoid plate is evident in the
large series of Protosalanx chinensis and Neo-
salanx j or dani I examined, even though these
are relatively small specimens. P. angusticeps is
based on a single specimen, the holotype, which
happens to be the largest referred to S. cuvieri
that I have examined. In vertebral counts and
other characters it apparently agrees well with
other material of S. cuvieri.

Salanx (Hemisalanx) prognathus (Regan, 1908b)
new combination

Hemisalanx prognathus REGAN, 1908b:445 (type-locality
Shanghai).

ISalanx brachyrostralis FANG, 1934a:257 (type-locality Nan-
king).

fReganisalanx brachyrostralis FANG, 1934b:509.

Metasalanx coreanus WAKJYA AND TAKAHASI, 1937 (otherwise
unpublished manuscript name placed in synonymy of H.
prognathus by WAK.IYA AND TAKAHASI 1937:293).

MATERIAL EXAMINED. — BMNH 1873.7.30.69, 111 mm,
Shanghai (holotype); CAS 51439, 34:97.5-120 mm, Kiangsu
Prov. (12:97.5-112 mm alcian-alizarin); CAS-SU 33990, 8:
94-1 14 mm, Tai Po, New Territory, Hong Kong.

This species has been recognized as repre-
senting a monotypic genus since its original de-
scription, and Wakiya and Takahasi (1937) even
placed it in a monotypic subfamily (Hemisalan-
ginae). Apart from its somewhat shorter, blunter
snout, and slight differences in dentition, how-
ever, it is very similar to specimens of the sub-
genus Salanx lacking a presymphyseal bone. The
magnitude of the differences between Hemisa-
lanx and Salanx (sensu stricto) is comparable to
that between the latter and Leucosoma, which I
also regard as only subgenerically distinct.

Regarding S. brachyrostralis, Wakiya and Ta-
kahasi (1937) placed it in the synonymy of S.
prognathus with a question mark, as is done here.

Neither Wakiya and Takahasi nor I have ex-
amined the holotype ("S. 4227, 144 mm total
length").

Wakiya and Takahasi (1937:293) placed Re-
ganisalanx normani in the synonymy of//, pro-
gnathus with a question mark, but I have ex-
amined the type-specimens and have reidentified
them as S. ariakensis.

Salanx (Leucosoma) reevesii (Gray, 1831)

1 Albula chinensis OSBECK, 1757 (type-locality West River at

Canton?; original not consulted).
Leucosoma reevesi GRAY, 1831:4 (type-locality China).
Leucosoma chinensis RICHARDSON, 1846:303.
Salanx reevesii VALENCIENNES in CUVIER AND VALENCIENNES,

1849:363, PI. 646.
Salanx chinensis GUNTHER, 1866:205.

MATERIAL EXAMINED.— AMNH 10336, 130 mm, Fukien;
AMNH 11161,74.5mm, Fukien; CAS-SU 1511,4:87.8-140
mm, Swatow (2:87.8-102 mm alcian-alizarin); CAS-SU 25738,
1 27 mm, Canton; CAS-SU 61189,153 mm, Castle Peak Bay?,
Hong Kong.

Salanx reevesi is second only to Protosalanx
chinensis as the largest and heaviest-bodied sa-
langoid. It is the only species of Salangidae in
which males with anal scales have not been re-
ported previously. A 130-mm specimen from
Swatow (CAS-SU 1511) is a mature male with
its anal fin greatly enlarged and 17 anal scales.
It is the only member of the subgenus Leuco-
soma, distinguished by a median row of 6-8 teeth
on the tongue or basihyal bone.

Albula chinensis Osbeck, 1765, has been iden-
tified with this species by various ichthyologists
following Richardson (1846) but, as pointed out
by Wakiya and Takahasi (1937:291-292), its
identity cannot be verified. While Albula chi-
nensis evidently is a member of the Salanginae,
there apparently are no type-specimen(s) extant
(Kullander, pers. commun., 1983) and it is im-
possible to tell from Osbeck's description which
species he had. The presence of teeth on the
tongue, which would positively identify it as S.
chinensis, is not mentioned.

In addition to having teeth on the tongue, Leu-
cosoma has jaw teeth that are somewhat larger
than those in the subgenera Hemisalanx and Sa-
lanx. In all other features of skeletal anatomy,
however, Leucosoma falls in the genus Salanx.
It has nearly the same vertebral counts as the
subgenus Hemisalanx, and its cranial shape ap-
pears to be identical with that in the subgenus
Salanx.

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

211

Salangichthyinae, new subfamily

Cranium only moderately depressed (as in
Sundasalangidae). Cranial fontanel with poste-
rior and apparently anterior portions open
throughout life. Head rounded anteriorly, pre-
maxillae failing to meet at midline, not project-
ing significantly beyond snout. Teeth greatly re-
duced in size, small or minute, those on maxilla
very numerous (about 1 5-40). Maxilla expand-
ed, much larger than premaxilla. Body moder-
ately elongate. Vertebrae 48-65 (66-79 in all oth-
er Salangidae).

This subfamily contains two genera, Neosa-
lanx and Salangichthys, formerly placed in Pro-
tosalanginae (Wakiya and Takahasi 1937).

Neosalanx Wakiya and Takahasi, 1937

Neosalanx WAKIYA AND TAKAHASI, 1937:282 (type-species, by
original designation, Neosalanx jordani WAKIYA AND TA-
KAHASI, 1937).

The species of this genus are poorly known. In
museum collections and older literature they are
usually misidentified as Protosalanx or Salanx.
Wakiya and Takahasi (1937) recognized four
species, three of which they described as new.
Although they recognized that Protosalanx an-
dersoni Rendahl belonged to Neosalanx, they
overlooked three previously described taxa, which
also apparently belong to it: Protosalanx brevi-
rostris Pellegrin, 1923; Protosalanx tangkahkeii
Wu, 1931; and Salanx argentea Lin, 1932. All
three of these were mistakenly placed by Fang
(1934a:240) in the synonymy of Protosalanx hy-
alocranius. I briefly examined the type-speci-
mens of Pellegrin's P. brevirostris in Paris, and
have since studied radiographs of them, but have
not seen the types of the taxa described by Wu
and Lin.

Wakiya and Takahasi (1937) distinguished four
species of Neosalanx, mainly on the basis of dif-
ferences in counts of vertebrae and fin-rays.
Without knowing more about interpopulational
meristic and other variation in Neosalanx, it is
difficult to evaluate the species. Specimens I ex-
amined tend to fall into groups, based on ver-
tebral counts, identical to those recognized as
species by Wakiya and Takahasi, and I have
identified my material accordingly.

Wakiya and Takahasi (1937) described Neo-
salanx as a new genus despite the fact that one
of its included species, Protosalanx andersoni

Rendahl, 1923, was proposed by Fang (1934a)
as type-species for his genus Paraprotosalanx. It
is obvious from Fang's account that he mistook
specimens of Protosalanx chinensis and proba-
bly either Neosalanx brevirostris or N. jordani as
Protosalanx andersoni and that his generic di-
agnosis is based mainly on P. chinensis. At this
point the nomenclaturally parsimonious solu-
tion would have been for Wakiya and Takahasi
to recognize Paraprotosalanx as a valid genus
with Protosalanx andersoni Rendahl, 1923 (not
of Fang 1934a), as its type-species. They chose,
however, to coin a new name and designated as
type-species the new species N. jordani. Given
that they did so, and that no publication has
appeared subsequently in which Paraprotosa-
lanx is treated except as a synonym of Neosa-
lanx, I provisionally recognize Neosalanx as val-
id. It should be noted, however, that judging from
Article 70 of the International Code of Zoological
Nomenclature ( 1 964 ed.) this case should be re-
ferred to the International Commission.

Neosalanx andersoni (Rendahl, 1923)

Protosalanx andersoni RENDAHL, 1923:92 (type-locality Chih-

li, Shan-Hai-Kuan).
Neosalanx andersoni WAKIYA AND TAKAHASI, 1937:285.

MATERIAL ExAMiNED.-NRM 10287, 79 mm, Chihli, Shan-
Hai-Kuan (holotype).

This species differs from all other Neosalanx
by its relatively large size, to 100 mm (vs. only
to 64 mm) and more numerous total vertebrae,
63-65 (vs. 59 or less). Wakiya and Takahasi re-
ported anal scale counts of 25-28 in TV. andersoni
and only 14-21 in other Neosalanx, but I find
only 21 or 22 anal scales on the male holotype.
N. andersoni is known only from rivers in Korea
and China flowing into the Yellow Sea.

The jaws of the holotype of Neosalanx ander-
soni appear to be almost entirely toothless. I de-
tected a single small conical tooth on the right
premaxilla, none on the left. The dentaries ap-
pear to be entirely toothless. The maxillae, al-
though having irregularities in the margin where
teeth may have been affixed, are toothless or al-
most toothless except for a very few minute teeth.
There seem to be no palatal or glossal teeth.

The anterior and posterior portions of the cra-
nial fontanel are widely separated and reduced
in size (especially the anterior portion). There is
no indication of a tectum medialis.

212

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

A small patch of minute, scaly breeding tu-
bercles occurs near the base of the 9th and 1 Oth
anal fin-rays, and more extensive areas of tu-
bercles have evidently been sloughed off. Pelvics
enlarged but without tubercles; lower jaw very
strongly projecting. Snout elongate, as in Sa-
langichthys (snout-tip to anterior margin of
orbit =4.2 mm).

Anal scale row continues onto ventral margin
of caudal peduncle a considerable distance (4
scales).

Tip of snout dorsally and tip of lower jaw,
dorsal surface of cranium over brain, dorsal sur-
face of body with irregularly distributed small
melanophores. No melanophores on anal fin, and
none or almost none on caudal fin, ventral body
surface, including pelvic and pectoral fin bases.

Neosalanx brevirostris (Pellegrin, 1923) new
combination

Protosalanx brevirostris PELLEGRIN, 1923:351 (type-locality
Tonkin).

? Protosalanx tangkahkeii Wu, 1931:219 (type-locality Amoy).

ISalanx argentea LIN, 1932 (type-localities Sangchang stream;
Henngchow Bay; Canton).

Protosalanx brevirostralis FANG, 1934a:232, 236, 240 (mis-
spelled; referred to synonymy of Protosalanx hyalocranius).

Neosalanx hubbsi WAKIYA AND TAKAHASI, 1937:284 (type-
locality Tien-tsin).

Neosalanx tangkahkeii taihuensis CHEN, 1954? (reference not
seen).

MATERIAL EXAMINED. -AMNH 10337, 22:43.8-60.7 mm,
Hunan (2:58.4-60.7 mm alizarin); AMNH 10480, 53.5 mm,
Tung Ting Lake, Hunan; AMNH 11155, 60.5 mm, Fukien;
AMNH 37044, 2:49.6-60.8 mm, Foochow, Fukien; AMNH
51690, 26:48-65 mm, Canton; CAS 52032, 2:46.1-48.3 mm,
Hong Kong (48.3 mm alizarin); CAS-SU 1540, 3:56.8-57.9
mm, Swatow; CAS-SU 68888, 3:56.0-58.2 mm, Seoul; MNHN
1922.184-189, 6:65-70 mm, Tonkin (syntypes of P. breviros-
tris); USNM 2 1 9923, 2: 5 1 .0-57.4 mm, Liang tsi Lake, Hupeh
(formerly identified as N. tangkahkeii taihuensis).

Protosalanx brevirostris has not been recog-
nized as a valid species or even referred to since
Fang (1934a), without examining the types, er-
roneously placed it as a junior synonym of Pro-
tosalanx hyalocranius. I briefly examined the
types, which are in rather poor condition, during
a visit to Paris in November, 1982, but did not
have specimens of other Neosalanx on hand for
comparison. I have since examined radiographs
of the types. Since their vertebral counts— 57(2),
58(2), 59(1)— correspond only to those reported
by Wakiya and Takahasi for Neosalanx hubbsi,
I conclude that they are conspecific.

Neosalanx brevirostris is known from the

mainland coast of Asia, from the Yellow Sea
coast of Korea south to Tonkin.

In most specimens of Neosalanx the premax-
illaries are toothless or have relatively few teeth
(1-6 reported by Wakiya and Takahasi (1937),
presumably based on examination of numerous
specimens, and 0-2 in a large number of speci-
mens I examined). Wakiya and Takahasi (1937)
reported only 1-2 premaxillary teeth in N. hubb-
si. Thus it is noteworthy that the 4 8. 3 -mm spec-
imen from Hong Kong (CAS 52032) here re-
ferred to this species has about 20 minute teeth
on each premaxilla. Tooth counts have not been
made on the type-specimens of N. brevirostris.

Despite considerable effort to trace its original
description, the taxon Neosalanx tangkahkeii
taihuensis is known to me only from an article
by Wang et al. (1980), an abstract of which ap-
peared in Aquatic Sciences and Fisheries Ab-
stracts. The article reports on its artificial fertil-
ization and larval development (see Addendum).

Neosalanx jordani Wakiya and Takahasi, 1937

Neosalanx jordani WAKIYA AND TAKAHASI, 1937:282 (type-
locality "River Rakuto, Corea").

MATERIAL EXAMINED. -AMNH 51705, 8:29-33 mm, Can-
ton; AMNH 51704, 47.5 mm, Foochow, Fukien; CAS 52028,
177:33.0-45.5 mm, Kiangsu, Chekiang Prov. (14:35.1-44.8
mm alcian-alizarin); CAS 52029, 1:36.8 mm, Hong Kong; CAS
52030, 31 1:22-58 mm, purchased in San Francisco; CAS-SU
68625, 160:35.3-47.1 mm, and UMMZ 55601, 20:35.5-41.3
mm, Pei-ho at Tien-Tsin (paratypes of Salanx hyalocranius
Abbott, 1901) (10:39.7-45.7 mm alcian-alizarin).

Wakiya and Takahasi (1937) characterize this
species as having 49-54 vertebrae (average 50.95).
The range of material I examined is 48-54. The
species is known from rivers along mainland coast
of Asia, from the Yellow Sea coast of Korea south
to Hong Kong.

Neosalanx reganius Wakiya and Takahasi, 1937

Neosalanx reganius WAKIYA AND TAKAHASI, 1937:283 (type-
locality "Ariake Bay, Kyushu, Japan," possibly erroneous).
Neosalanx regani MATSUBARA, 1955:214 (unjustified spelling).

Wakiya and Takahasi (1 937:283) reported 52-
56 (average 53.75) vertebrae for this species. It
is known only from the type-specimens, sup-
posedly collected in Ariake Bay. I have not ex-
amined any Neosalanx from Japan.

Salangichthys Bleeker, 1860

Salangichthys BLEEKER, 1860:101 (type-species, by monotypy,
Salangichthys microdon BLEEKER, 1 860).

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

213

Salangichthys microdon Bleeker, 1860:101

Salangichthys microdon BLEEKER, 1860:101 (type-locality

Yeddo-Tokyo).
Salangichthys kishinouyei WAKIYA AND TAKAHASI, 1913:552

(type-locality Seishin, Korea and Shimane-ken, Japan).

MATERIAL EXAMINED. -AMNH 13 149, 28:50-64 mm, Dao-
mori-ken, Japan; CAS 52033, 6:74.8-86.7 mm, Kawasaki; CAS
52034, 9:31.6-36.1 mm, Lake Kituara (alcian-alizarin); CAS-
SU 134, 10:78-92 mm, Tokyo (5:81-86 mm alcian-alizarin);
CAS-SU 22637, 61:32-67 mm, Sendai, Matsushima Bay, Ja-
pan.

Salangichthys kishinouyei originally was dis-
tinguished from S. microdon by Wakiya and Ta-
kahasi (1913) on the basis of its having 1) man-
dibular, maxillary, and palatal teeth more widely
spaced; 2) snout shorter; 3) anal fin origin in
females more anterior; and 4) threads of external
egg membrane thicker, their ends club-shaped
and ending freely without fusing to each other to
form a ring. They later concluded that these dif-
ferences represent intraspecific variation in pop-
ulations of 5. microdon on the Japan Sea coast
of Sakhalin, Korea, Honshu, and Kyushu and
placed S. kishinouyei as a synonym of S. micro-
don (Wakiya and Takahasi 1937:279-280).

Matsuoka and Iwai (1983, Fig. 2) illustrated
an alcian-alizarin preparation of S. microdon (lo-
cality not indicated) with 65 total vertebrae. The
highest number of vertebrae otherwise known in
Salangichthys is 63 (Table 2).

Salangichthys ishikawae Wakiya and
Takahasi, 1913

Salangichthys ishikawae WAKIYA AND TAKAHASI, 1913: 552
(type-locality Miyagi-ken, Japan).

MATERIAL EXAMINED. -CAS-SU 6780, 9:67.6-74.3 mm,
Same, Rikuoku (3:70.3-74.2 mm alcian-alizarin); CAS-SU
68878, 20:48-63 mm, Pacific coast of Aomori-ken, northern
Honshu; CAS-SU 68889, 70.7 mm, Ariake Bay, Kyushu (lo-
cality possibly erroneous).

Salangichthys ishikawae is distinguished from
S. microdon mainly by having 20-27 pectoral
fin-rays instead of only 14-19 (see Senta 1973c).
Differences in counts of vertebrae and anal scales,
based on few specimens, are of doubtful signif-
icance (see Table 2).

According to Wakiya and Takahasi (1937:281),
S. ishikawae "lives in water of rather higher sa-
linity than S. microdon . . . and seems to be a
northern species," known only on the Pacific coast
of Honshu. The specimen herein reported from
Ariake Bay, Kyushu, represents a southerly range
extension of about 500 km if its locality data are

correct. It was found with two specimens of Sa-
lanx ariakensis collected by Kishinouye in Ar-
iake Bay (CAS-SU 8574).

Sundasalangidae Roberts, 1981

Primary pectoral girdle consisting of single
median element (identical to the condition in
various larval teleosts, but unknown in adults of
any other teleost). Pectoral fin rayless, supported
by a single undivided basal radial. Pelvic fin with
5 rays. Each half of pelvic girdle with two para-
pelvic cartilages (unknown in any other teleosts).
Adipose fin absent. Adult males without anal
scales or sexually dimorphic anal fins. Neural
spines much more elongate than in Salangidae.
Hemal arches of last 1 2 or so abdominal verte-
brae with elongate hemal spines (abdominal ver-
tebrae without hemal spines in all other salan-
goids). Vertebrae 37-43. Caudal fin skeleton with
parhypural fused to hypurals 1 and 2 (parhypural
separate in all other salangoids).

If specialized is defined as deviation from the
morphology of any known group of salmoni-
forms that could possibly serve as the ancestral
stock of Salangoidea, then Sundasalanx is by far
the most specialized of all salangoids.

The samples of Sundasalanx available at the
time of their original description differed so much
(in regard to eye size, non-overlapping vertebral
counts, etc.) that it seemed to me that they could
not be the same species. Samples examined sub-
sequently, from Muar and the Mekong, tend to
be intermediate between the two described
species. I have tentatively identified the new
samples with Sundasalanx praecox, but the
question of the number of species in the genus
should be reconsidered when more material be-
comes available. The genus has not been found
yet in Sumatra but is to be expected there.

Sundasalanx Roberts, 1981

Sundasalanx ROBERTS, 1981:297 (type-species, by original
designation, Sundasalanx praecox ROBERTS, 1981).

Sundasalanx microps Roberts, 1981

Sundasalanx microps ROBERTS, 1981:300 (type-locality Ka-
puas River at Kampong Nibung, about 100 km NE of Sin-
tang and 7 km NE of Selimbau).

MATERIAL EXAMINED.-CAS 44220, paratypes, 34:14.6-19.9,
Kapuas River at Kampong Nibung (paratypes) (7:13.5-18.0
mm alcian-alizarin).

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

Sundasalanx praecox Roberts, 1981

Sundasalanx praecox ROBERTS, 1981:299 (type-locality Khlong
Falamee, Tale Sap, about 2 km W of Pak Payoon, Kra Isth-
mus, Southern Thailand).

MATERIAL EXAMINED. -CAS 5203 1 , 6: 1 6.4-1 7.7 mm Khlong
Falamee, Tale Sap, near Songkhla, Thailand (paratypes; alcian-
alizarin); UMMZ uncat. and CAS 52036, 65:15.7-22.5 mm,
Mekong River and tributaries in Thailand from Ban Dan to
Nakon Phanom (4:18.2-20.3 mm alcian-alizarin; USNM
229304 and CAS 52037, 121:12.3-18.0 mm, Muar River 7-
1 8 miles inland from Bandar Maharani (formerly Muar City)
(12:16.5-17.8 mm alcian-alizarin).

When more material becomes available and
the species of Sundasalanx are re-examined, the
pigmentation of the Muar specimens (CAS 52037,
USNM 229304) should be taken into consider-
ation. In the Sundasalanx examined, each sam-
ple tends to have a more or less distinctive pig-
mentation or pattern of melanophore distribution
shared by all of the specimens. Thus the type-
series of S. microps is characterized by having a
series of large mid ventral melanophores, one per
myotome, extending from the pectoral fin to the
anal fin. Midventral melanophores are absent in
the type-series of S. praecox. In the Muar spec-
imens here identified as 5". praecox, however, a
series of midventral melanophores extends from
about pelvic-fin origin to the anal fin, thus re-
sembling the pigmentation of S. microps. On the
other hand, each of the Muar specimens has a
series of large melanohores along the base of
the anal fin, one melanophore between the base
of each branched fin-ray. Other samples of Sun-
dasalanx examined do not exhibit this feature.

In the original description of S. praecox (Rob-
erts 1 98 1 :299) it is stated that the type-series has
maxillary teeth about 15-19 vs. about 30 in S.
microps; the reverse is true.

DISCUSSION

Anyone who has looked into recent accounts
of salmoniform classification realizes that it is in
disarray. In this group current classifications are
based largely on skeletal anatomy, and as long
as the skeletal anatomy of major groups such as
salangoids remain uninvestigated and others only
partially investigated, no stable classification can
be expected. McDowall (1969) briefly examined
the skeletal anatomy of "Salangichthys micro-
don" (actually Neosalanxl) stained with alizarin
and concluded that Salangidae "is not part of the
galaxioid radiation." He also stated "a more

complete study (of salangid skeletal anatomy)
may be desirable, should a full range of material
be available, especially if it should reveal some
species with more complete ossification to enable
more precise determination of affinities." A
species with somewhat more complete ossifica-
tion actually does exist— Protosalanx chinen-
5/5— but even in this species truly adequate ob-
servations of skeletal anatomy cannot be based
on specimens stained solely with alizarin. The
question is no longer relevant, however, since
adequate skeletal preparations of salangoids usu-
ally can be obtained using the alcian-alizarin
technique.

All modern accounts of salangid classification
agree in placing them in the order Salmoni-
formes. My information on salangoid skeletal
anatomy, however, has not provided me with
obvious answers about their relationships to oth-
er salmoniforms, but has only emphasized their
distinctness. The presence of two (marginal or
submarginal) rows of teeth on the basihyal tooth-
plate of Protosalanx confirms the integrity of the
order Salmoniformes and the placement of Sa-
langoidea within this order but nothing more.
The presence of a taenia medialis in the cranium
of young Protosalanx and a single specimen of
Neosalanx suggests a shared derived character
(synapomorphy) with osmeroids, but this char-
acter may well prove primitive for salmoni-
forms, perhaps to be found in many of them.

Higher classification of Salmoniformes has
been the subject of considerable interest in the
last two decades or so, with contributions by
Gosline (1960), Greenwood et al. (1966), Weitz-
man (1967), McDowall (1969), Rosen (1974),
Klyukanov (1975), and Fink and Weitzman
(1982). A major issue is whether esocoids (pikes,
northern mud-minnows, and relatives) are Sal-
moniformes, and should include the southern
Lepidogalaxias, as advocated by Rosen. This
view is contested by Fink and Weitzman, who
exclude esocoids from Salmoniformes and relate
Lepidogalaxias to Galaxiidae and osmeroids.
Another major issue is whether relations of the
southern "salmonoids" (Galaxiidae, etc.) lie with
the northern salmonoids (Salmonidae, etc.) or
with osmeroids. The hypothesis that stomiatoids
are Salmoniformes (Weitzman 1967) has been
rejected by Rosen ( 1 974), Klyukanov (1975), and
Fink and Weitzman (1982). Fink and Weitzman
also point out that no satisfactory evidence has

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

215

yet been advanced to support the hypotheses of
relationships among osmeroids, salmonids, and
galaxioids. Here I shall comment briefly on re-
lationships of the salangoids to other Salmoni-
formes, particularly osmeroids and galaxioids,
and present some evidence bearing on relation-
ships of Lepidogalaxias.

Gosline (1960) suggested a close relationship
of Osmeridae, Salangidae, and Plecoglossidae,
distinguishing them from the Southern Hemi-
sphere Retropinnidae, Aplochitonidae, and Ga-
laxiidae. Greenwood et al. (1966) placed Salan-
gidae in a suborder Galaxioidei including
Retropinnidae, Galaxiidae, and Aplochitonidae
without explanation. McDowall (1969) conclud-
ed that Salangidae are a very specialized offshoot
of the salmonoids and not part of the galaxioid
radiation.

In observing the skeletal anatomy of salan-
goids I have been watchful for specialized char-
acters indicative of phyletic relationship to other
Salmoniformes. While my study has revealed
highly specialized characters (such as the hyopal-
atine) indicating monophyly of salangoids, it has
not provided (or at least I have not noticed) char-
acters that would link salangoids in a monophy-
letic taxon with Osmeridae or any other group.
Rosen (1974) pointed to specializations of the
anal fin and associated scales in males of Lepi-
dogalaxias and Mallotus, but concluded (p. 304)
that these do not indicate relationship to Sa-
langidae, and I agree. The anal scales and anal
fin modifications of male salangoids appear to
be a unique specialization, as does the hyopal-
atine. Salangoids lack some skeletal features
found in Osmeridae, notably in the ethmoid re-
gion. Whether this absence is due to loss or re-
flects a primitive condition is unclear. Devel-
opment of a tectum taenia medialis in the cranial
fontanel, characteristic of the osmeroid chon-
drocranium, occurs in some salangoids and may
be indicative of relationship, but further obser-
vation may reveal that the character is wide-
spread in Salmoniformes. The peculiar mor-
phology of the salangoid egg case (Wakiya and
Takahasi 1937, Okada 1960) may be similar to
that in osmeroids (compare photographs of Hy-
pomesus olidus and Salangichthys microdon eggs
in Chyung 1961, figs. 242-243). Similar spe-
cializations possibly also occur in Plecoglossus
but have not been reported in any of the northern
salmonoids or in galaxioids.

According to McAllister (1963:6) "the neo-
tenic Salangidae strongly resemble larval Os-
meridae but may be distinguished by their re-
duced pointed head and elongated anterior
portion of the body, as well as osteological
characters." Skeletal preparations of larval Os-
meridae examined by me differ from salangoids
in many respects. There are certain similarities
in appearance of the largely transparent and lightly
pigmented larvae of osmeroids and salangoids
but these are mainly such as are to be found in
larvae of non-related teleosts, and their value in
assessing relationships is dubious. Skeletal anat-
omy of larval osmeroids I have examined differs
from that of salangoids almost as much as does
skeletal anatomy of adult osmeroids. I have not
examined skeletal anatomy of Plecoglossidae, but
this family appears to be very specialized and
there is no evidence that it is particularly closely
related to Salangidae. Gosline (1960:346) and
others have mentioned certain similarities be-
tween the salangids and the extraordinarily spe-
cialized galaxoid Lovettia but I believe such re-
semblance is due to independently acquired
neotenic characters.

Considerable interest has centered on the phy-
logenetic significance of the little salmoniform
Lepidogalaxias salamandroides recently discov-
ered in western Australia (Mees 1961). Among
many peculiar features, it has in sexually mature
males an extraordinarily modified anal fin par-
tially covered by a sheath of anal scales sugges-
tive of the anal scales of male Salangidae. The
extremely complex modifications of the anal fin-
rays go far beyond that seen in the anal fin of the
salangids or any other salmoniform, and I doubt
that in the relatively simple modifications of the
anal fin in male salangids any uniquely shared
specializations (or synapomorphies) with Lepi-
dogalaxias can be recognized. The presence of a
sheath of enlarged anal scales, on the other hand,
demands closer comparison with those of sa-
langids, which are otherwise unique among sal-
moniforms (and perhaps all other teleosts). The
anal scales in Lepidogalaxias are greatly enlarged
and disposed in two main rows, rather than a
single row, as in salangids. Two 3 1-34-mm spec-
imens I examined have seven to eight scales in
the upper row and three in the lower. There also
appear to be some scales or scalelike structures
associated with the vent itself. Unlike that of
salangids, however, the anal sheath covers the

216

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

mesopterygoid

ectopterygoid
palatine
premaxilla

metapterygoid

symplectic
interhyal

hyomandibula
opercle

maxilla
dentary

articular

angular
quadrate

subopercle

1 mm

preopercle

interopercle

FIGURE 22. Lepidogalaxias salamandroides (uncat., 23.5 mm). Jaws, facial ones, and suspensorium (lateral view).

anal fin, especially the modified portions. In sa-
langids the anal scales are on the body above the
anal fin, and the rays themselves are entirely ex-
posed. Whatever the phyletic significance, it
would certainly be of interest to know more about
the functional significance of the anal scales and
modified anal fins in Lepidogalaxias and in Sa-
langidae.

Fink and Weitzman (1982) suggested that a
single row of mesopterygoid teeth is a synapo-
morphy indicating monophyly for the osmeroids
(including Salangidae) and galaxioids (including
Lepidogalaxias). Although Gosline ( 1 960) stated
that the mesopterygoid is absent in Salangidae,
it is actually present in most of them. But while
most Salangidae have palatal teeth, the salangoid
mesopterygoid is invariably toothless, even in
Protosalanx and Salanx chinensis which have
well-developed basihyal teeth. Thus the concept
of a "tongue-bite" character, based on basihyal
and mesopterygoid teeth, and uniting galaxioids
and osmeroids, does not hold for salangoids.

While my observations of salangoid skeletal
anatomy have not provided me with the key to
their higher relationships, I earnestly hope that
they may do so for future workers who are able
to make more extensive comparisons. In ex-
amining Lepidogalaxias I find no characters in-
dicative of close relationship to Osmeridae or

Salangidae, but the structure of the jaws and gill
cover reveals specialized characters linking this
strange western Australian fish to the southern
galaxioid radiation.

Based largely on the assumption that Lepi-
dogalaxias possesses uniserial mesopterygoid
teeth, and without having examined skeletal ma-
terial, Fink and Weitzman (1982) suggested its
relationships lie with osmeroids and galaxioids.
I have examined skeletal anatomy of several lar-
val and juvenile or subadult Lepidogalaxias; the
largest of these has a well-developed median patch
of teeth on the prevomer and a pair of well-
developed tooth patches on the anterior palatine
area (ectopterygoid? = endopterygoid of Mees
1960) but the large, well-developed mesoptery-
goid is toothless. On the other hand, specializa-
tion of bones in the jaws and gill cover (Fig. 22)
indicates relationship with galaxioids. The ex-
traordinary fimbriate condition of the bony oper-
cle, also of the subopercle, appears to be a highly
specialized character shared only (i.e., synapo-
morphic) with galaxioids (see McDowall 1969,
Fig. 3b—e, and Rosen 1974, Fig. 13). The short
and laterally directed premaxilla and toothless
maxilla with its strongly concave margin may
represent another synapomorphy with galax-
ioids. My observations suggest that Lepidoga-
laxias is indeed related to Galaxiidae, as sug-

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

217

gested by Nelson (1970), but I doubt that it
represents the "primitive sister group ofgalaxiids
or galaxioids"; it is more likely to be a highly
specialized galaxiid or galaxioid derivative. As
pointed out by Scott (1966), its osteology should
be examined carefully and thoroughly; this has
yet to be done. For the present I would like to
point out a difference between the branchial
arches of Lepidogalaxias and Umbridae which
merits further investigation. In Lepidogalaxias
and Umbridae the basibranchial plate has five
basibranchials, and the fifth is cartilaginous. In
Lepidogalaxias the basibranchial plate is appar-
ently immobile; that is, the basibranchials ap-
parently do not move backward and forward in
relation to each other. In Umbridae, however,
as exemplified by Novumbra hubbsi, basibran-
chial 5 is movably articulated to basibranchial 4
in such a fashion that it can be rocked back and
forth beneath it; and thus the pair of toothplate-
bearing fifth ceratobranchials, firmly attached to
basibranchial 5, are also moved back and forth.
Whether such basibranchial mobility occurs in
other Umbridae or in esocoids generally is un-
known; it has not been observed in galaxioids,
osmeroids, or salangoids (the latter apparently
lack basibranchial 5).

In assessing phylogenetic relationships one
should not be overly impressed by the presence
of primitive characters, even in groups in which
such characters supposedly have been lost for a
long time.

Atavism, the expression of ancient characters
"buried in the genome," occurs far more often
than generally recognized. This, rather than a
Lamarckian interpretation, is doubtless the cor-
rect explanation for the appearance of breeding
tubercles on the palms of the midwife toads (Al-
ytes obstetricans) painstakingly studied by Kam-
merer (see Koestler 1973). Characters such as
eyes, teeth, scales, bone, or the pelvic girdle may
be repeatedly suppressed, and may not be phys-
ically expressed in any members of quite large
groups, without ever having been lost from the
genome. For purposes of phylogenetic analysis,
I suggest it is parsimonious to assume that prim-
itive characters are never lost from the genome,
and that this is really why it is futile to rely on
them. Thus the fully scaled condition of Lepi-
dogalaxias does not suggest to me that it rep-
resents the primitive sister group of the otherwise
scaleless galaxioids.

I believe that utilization of the caudal skeleton
as a guide to phyletic relationships among te-
leosts, especially those with the primitive teleost
complement of six separate hypurals and 10 + 9
principal caudal fin-rays, has inevitably resulted
in confusion of primitive with advanced char-
acteristics in the caudal fin skeleton. While te-
leosts as a whole exhibit great diversity in their
caudal skeleton (Monod 1 968), the main features
of the caudal skeleton are remarkably similar in
many teleosts with forked caudal fins and the
primitive complement of principal caudal fin-
rays. Thus the caudal fin skeleton of salangoids
is strikingly similar in many respects to that of
Flops, many clupeoids, characoids, and cypri-
noids as well as of osmeroids and other salmon-
iforms. There are two possible explanations (or
hypotheses) for such similarities, both predicated
upon the assumption that the caudal fin structure
of Elops and the others is primitive for teleosts.
The first, and traditional, explanation is that all
descended from ancestors that never deviated
from morphologically primitive caudal fins. The
second is that teleosts with secondarily general-
ized caudal fins have repeatedly reverted to a
primitive type of caudal fin skeletal morphology.

Salangoids often have been referred to as neo-
tenic. According to Jordan and Snyder (1902:
592), "the straight alimentary canal, distinct
muscle segmentation, very thin ventral wall of
the abdominal cavity, and other characters of
salangoids suggest a larval stage of develop-
ment." Among other characters, they were cer-
tainly thinking of the near perfectly transparent
state of the fish in life. Thus when the fish die,
the flesh turns perfectly white, and, in the absence
of scales, the myotomes stand out very clearly,
as in many larval fish. Berg (1947), Gosline
( 1 960), Weitzman (1967), and McDowall (1969),
remarked that Salangidae seem to be neotenic
but did not elaborate. If the term neotenic means
simply attaining sexual maturity while retaining
some larval characteristics, salangoids are surely
neotenic, probably more so than any other sal-
moniform fishes. The following characteristic
features of salangoids are among those which
may be neotenic:

1) Body almost entirely transparent in life.

2) Pectoral fin pedunculate, with a broad, fleshy,
pedestallike base more or less free from the lat-
eral musculature of the body, and placed high on

218

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 13

the side of the body, just like the pedunculate
pectoral fin of many lower teleost larvae.

3) Skeleton in large measure cartilaginous;
many bones found in adults of other salmoni-
forms absent.

4) Body scaleless throughout life except for
anal scales in sexually mature male salangids.

5) Ventral body musculature incomplete, ev-
idently due to an arrested ventral myotomic pro-
gression, so that the ventral abdominal wall is
thin and non-muscular.

6) Left and right halves of pelvic girdle more
or less widely separated from each other, failing
to form a cartilaginous union by means of pos-
teromedian processes as in most other salmon-
iforms.

7) Abdomen posterior to pelvic fin with a
membranous median keel.

8) Fourth gill arch with well-developed hy-
pobranchials.

9) Primary pectoral girdle consisting of a me-
dian scapulocoracoid in Sundasalangidae (a con-
dition also present in larvae of Elops and other
lower teleosts).

10) Dorsal and anal fins placed relatively far
posteriorly (least so in Protosalanx).

On the other hand, salangoids exhibit consid-
erable diversification and a number of peculiar
modifications or specializations which are ob-
viously not neotenic, including:

1) Marked sexual dimorphism in Salangidae,
involving enlarged pectoral and anal fins, mod-
ified anal fin-rays, and development of the anal
scales in sexually mature male salangids.

2) Strongly depressed cranium and skull;
marked ventromedian curvature of maxilla.

3) Voracious feeding habits and canine den-
tition, especially in Salanginae.

4) Vertebral counts of 37-79, the highest counts
found in Salanginae with excessively elongate
body form.

5) Proximal radials of pectoral fin highly mod-
ified in all Salangidae; pectoral fin-rays very nu-
merous in all Salangidae except Salanginae.

6) Fusion of cartilaginous hyomandibular and
mandibular arches to form a hyopalatine.

7) Fusion of basibranchials and hypobranchi-
als in gill arches of Sundasalangidae.

Thus salangoids resemble larval fish in many
ways. But the problem arises of distinguishing
between characters that are truly neotenic and
characters that represent convergence of adults
with larvae. Upon first observing the beautifully

simple condition of the primary pectoral girdle
in Sundasalanx, and taking into consideration
the fused condition of ventral elements in the
branchial arches of Sundasalanx and of the
pterygoquadrate and hyomandibula in all sa-
langoids, I was inclined to regard it as due to
secondarily evolved simplification and/or re-
duction and fusion of the primitively separate
left and right halves. But a morphologically iden-
tical "median" pectoral girdle and "fused" radial
plate was reported in larval clupeoids by Good-
rich (1922) and I have found it in larval Elops
hawaiiensis (Fig. 19b) and anchovies. In all of
these larvae, as in Sundasalanx, the scapulocor-
acoid has three clearly divided portions (a trans-
verse median bar, an ascending process, and a
posterior projection) and the basal radial or plate
is perforated by three foramina. In further de-
velopment, the three foramina of the pectoral
plate presumably enlarge until the basal plate in
all of these forms except Sundasalanx divides
into proximal radials 2-5. (The primitive num-
ber of proximal radials for all teleosts appears to
be 5, observed even in many teleosts having
highly modified pectoral fins.) Sundasalanx is
the only known fish which retains a median scap-
ulocoracoid at sexual maturity. The other salan-
goids presumably have such a median pectoral
girdle as larvae, but although the scapulocora-
coid is apparently cartilaginous in all Salangidae
examined, it is clearly divided into two halves
in all specimens examined, including Salangich-
thys microdon of only 39 mm SL. Goodrich
(1922:508) tentatively concluded that the fusion
of the endoskeletal pectoral girdle he found in
young clupeids represents a specialization pe-
culiar to larval Clupeidae. The present finding
that morphologically identical girdles occur in at
least some Elopomorpha and Salmoniformes as
well suggests that it is indeed a primitive char-
acteristic of teleosts. The duration of the median
condition apparently corresponds more or less
closely with a period when the pedunculate pec-
toral fins are being used most actively, the pri-
mary pectoral girdle is still largely or entirely
cartilaginous, and the mesocoracoid arch has not
developed.

ADDENDUM

The galley proofs of this paper were already
set when Prof. Xin-Luo Chu of the Kunming
Institute of Zoology of Academia Sinica visited
the California Academy of Sciences (January

ROBERTS: SALMONIFORM SUPERFAMILY SALANGOIDEA

219

1984) bringing specimens of Neosalanx tangah-
keii taihuensis for me to examine. These speci-
mens, CAS 54330, 4: 69.7-76.8 mm, were caught
in Kunming Lake where the species was intro-
duced in 198 1 from artificially reared stock orig-
inating in Lake Taihu. The specimens have total
vertebrae 56(1), 57(2), 59(1) and gill rakers 15(2),
16(1), 18(1). I therefore conclude that N. tan-
gahkeii taihuensis is a junior subjective synonym
of N. brevirostris. N. brevirostris is a valuable
commercial fish and much of the production (es-
pecially of Lake Taihu) is marketed abroad. In
the near future it may be widely introduced in
lakes in China which lie outside the natural range
of Neosalanx.

It may be worthwhile for Chinese workers to
investigate the aquacultural potential of the other
two species of Neosalanx occurring in China. N.
andersoni, which may grow slightly larger than
TV. brevirostris, has a relatively restricted north-
erly distribution, indicating that it is adapted to
colder waters. N. jordani, the smallest species,
has a wide range largely overlapping that of N.
brevirostris, but my observations indicate that
the two species usually do not occur together in
nature. N. jordani has fewer gill rakers and this
together with its smaller size indicates a probable
difference in feeding habits. It could be of prac-
tical as well as scientific value to compare the
ecology and fisheries biology of these three
species.

ACKNOWLEDGMENTS

For providing specimens, information, advice,
or assistance I wish to thank the following per-
sons: Tokiharu Abe, Janine Abel, Gerald R. Al-
len, Vladimir Barsukov, Marie-Louise Bauchot,
W. H. Butler, Xin-Luo Chu, Alexandra Creigh-
ton, Oliver Crimmen, Robert Drewes, Norma
Feinberg, Bo Fernholm, W. I. Follett, Michael
Hearne, Susan Jewett, Sven Kullander, Robert
McDowall, Gareth Nelson, Han Nijssen, Nicolai
Parin, Ingrid Radkey, Walter Rainboth, Donn
E. Rosen, Lars Wallin, Stanley H. Weitzman,
and Alwyne Wheeler.

This study was carried out mainly in the De-
partment of Ichthyology of the California Acad-
emy of Sciences, National Marine Fisheries Lab-
oratory at Tiburon, and the Tiburon Center for
Environmental Studies (of San Francisco State
University), and was supported in part by grant
DEB77-24574 in the Systematic Biology Pro-
gram of the National Science Foundation.

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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 14, pp. 221-238, 17 figs., 1 table July 12, 1984

By
Timothy C. Tricas

Department of Zoology, University of Hawaii at Manoa,
Honolulu, Hawaii 96822

and
John E. McCosker

Steinhart Aquarium, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118

ABSTRACT: The feeding behavior of the white shark (Carcharodon carcharias) was studied at Dangerous
Reef, South Australia. Cinematographic analyses of shark feeding patterns show that a single bite action is
comprised of a uniform sequence of jaw and head movements. The components are: 1) snout lift, 2) lower-
jaw depression, 3) palatoquadrate protrusion, 4) lower-jaw elevation, and 5) a bout-ending snout drop. Du-
rations for a complete bite action ranged from 0.750 to 1.708 s (x = 0.985 s) for a 3.5 m (TL) subject. Various
approach behaviors to baits were also documented.

The stomach contents of nine white sharks captured in northern and central California waters consisted
entirely of fish prey associated with inshore and pelagic habitats. Records of the stomach contents of 24
additional sharks were combined and analyzed, and indicated fish to be the most frequent prey items, while
marine mammals were also common. Analysis of prey type in relation to shark size shows small sharks (<3
m) feed primarily on fish prey, while larger sharks feed on marine mammals, especially pinnipeds.

Cursory field experiments and observations indicate sharks detect and are attracted to electric fields.
Telemetric studies of white shark thermal biology show that they are warm-bodied, approximately 4-5°C
above ambient water temperature.

Length-weight records for 127 sharks were analyzed and found to have the relationship: W = 3.8 x 10~6
L3 1S, where W is weight in kg and L is length in cm. The largest reliable record for a white shark is that of
a 6.4-ni, 3324-kg specimen captured near Cojimar, Cuba, in 1945.

A hypothesis is proposed to explain the "bite and spit" paradox related to attacks on pinnipeds and humans.
Comments concerning the risk associated with contemporary surfboard design are included.

INTRODUCTION Collier 1964; Follet 1974; Ellis 1975; McCosker

The white shark (Carcharodon carcharias) (Fig. 1 98 1). It is circumglobal in distribution, but most

1) is the largest piscivorous marine fish in the commonly inhabits the coastal temperate waters

world and is well known for its aggressive be- of North America, South Africa, and South Aus-

havior and potential threat to humans (Fast 1955; tralia.

[221]

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14

FIGURE 1. Tagged male white shark swimming near the surface at Dangerous Reef, South Australia. Photo by Al Giddings.

In spite of its size and fearsome reputation,
surprisingly little is known about the natural his-
tory and behavior of this large fish. Almost all
published information on the general biology of
white sharks comes from anecdotal observations
and notes obtained from commercial fishing or
whaling operations (Squire 1 967), regional species
lists and range extensions (Bigelow and Schroe-
der 1948; Day and Fisher 1954; Royce 1963),
and newspaper articles on captures by fishermen
or accounts of attacks on humans.

The predatory behavior and feeding mechan-
ics involved in prey capture by white sharks has
remained, until the recent application of scuba
and high speed photography, essentially un-
known. Previous studies on the feeding mor-
phology of other species of sharks were based
largely on anatomical data where muscle and
supportive tissue functions were inferred from
examination of preserved specimens (Luther
1909; Haller 1926). This approach provided
functional insight, based largely on articulations
and spatial arrangements of skeletal tissues and
head musculature. In some cases, however, the
inflexibility of preserved materials has led to
misinterpretations of the true mechanics of jaw
protrusion and feeding in sharks (see Compagno

1977). Whereas examination of fresh pliant spec-
imens may be more appropriate for functional
analyses, they still provide only speculative data
on sequential and temporal relationships of
structures involved in feeding activity. Moss
(1972) provided a qualitative analysis of feeding
mechanisms in living carcharhinid sharks using
observational, photographic, and electrical mus-
cle stimulation techniques. Studies on the tem-
poral and sequential mechanics of feeding be-
havior in sharks are still lacking, however, when
compared to the more thorough cinematograph-
ic studies on teleostean fishes (Osse 1969; Liem
1978; Lauder 1980).

Because white sharks are rarely captured, doc-
umentation of their food habits is scattered
throughout the literature. Most records come
from notes on the stomach contents of dead fish
(Schroeder 1938; Bonham 1942; LeMier 1951;
Scattergood 1962) or from fortuitous observa-
tions of feeding in the field (Day and Fisher 1954;
Pratt et al. 1982). More complete accounts are
provided on the relationships of white sharks to
pinnipeds (Ainley et al. 1981; and Le Boeuf et
al. 1982) and sea otters (Ames and Morejohn
1 980). There still remains, however, the need for
a comprehensive collation of the prey items tak-

TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK

223

en by this predator so that a more complete as-
sessment of predator-prey relationships can be
made.

This paper presents new data obtained during
a recent expedition to South Australia that relate
to white shark predatory behavior and general
biology. We analyze the feeding mechanics of
white sharks in the field by use of cinemato-
graphic techniques and provide information on
their sensory biology and thermal physiology. In
addition, we have synthesized previously pub-
lished and unpublished data on the length-weight
relationships, predator-prey interactions, and
general behavior of this shark. Based upon what
is known about the predatory behavior of white
sharks, we present a new interpretation of the
curious pattern of non-feeding attacks upon ma-
rine mammals and humans.

STUDY AREA AND METHODS

White sharks were studied in the field during
a ten-day period in January 1980, in waters near
Dangerous Reef, South Australia. The reef con-
sists of two small, low islands approximately 16
km east of Port Lincoln (Fig. 2). Sharks were
attracted to the 20-m vessel, Nenad, using tuna
and meat byproducts as bait. Sharks feeding on
baits both at and below the surface were pho-
tographed using Actionmaster 500 cameras and
7247 Kodak color reversal film exposed at shut-
ter speeds of 24 and 200 frames per second.
Frame-by-frame analyses were performed on a
Movieola 16-mm film editor.

Stomach content and morphological data from
nine sharks on record at the California Academy
of Sciences were analyzed. These data were then
combined with other published records and fur-
ther examined. To prevent multiple entries of a
record into the analyses, only well-documented
reports that included information on capture lo-
cality, number of sharks sampled, measured
lengths and weights (no estimations), and specific
prey types were used.

Two types of ultrasonic telemetry packages,
constructed by the senior author, were used in
this study to monitor shark body and ambient
water temperatures. All transmitter circuits con-
sisted of a crystal-controlled oscillator (carrier
frequencies = 31.700 or 32.768 kHz) gaited by
a thermistor-controlled pulse circuit sensitive
from 10°C to 33°C. The first tag consisted of a
single transmitter with a thermistor probe

FIGURE 2. The study area, Dangerous Reef, South Austra-

lia.

(embedded on the surface of the transmitter
housing) that monitored ambient water temper-
ature around the animal. Its dimensions were
4.6 x 3.2 x 2.0 cm, and it weighed approxi-
mately 60 g in air. The second unit consisted of
a cylindrical package with two transmitters of
different carrier frequencies. One transmitted
temperature data from a thermistor in contact
with the surrounding water, the other from a
thermistor embedded under the barb of a dart at
the end of a 3 1 cm-long wire leader. Total pack-
age dimensions were length 17 cm x diam. 3.2
cm, with a weight of approximately 100 g in air.
The water temperature sensing package was
applied from underwater using scuba (and a cage).
A stainless steel dart was attached to an appli-
cator tip on the end of a speargun shaft, and shot
3 cm deep into the shark's mid-lateral muscu-
lature. The dual-temperature sensor package was
applied externally to another shark from the
swimstep of the research vessel via barb and ap-
plicator pole. Signals were tracked with a tune-
able ultrasonic receiver and a staff-mounted di-
rectional hydrophone. Absolute maximum range
of the transmitter-hydrophone system under ide-

224

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14

100 -

50 -

In W = -12.47 + 3.15 (In L)

Total Length (cm)

FIGURE 3. Length-weight relationship for the white shark.
Data taken from complete records for 127 sharks. Functional
(geometric mean) regression equation given on figure (see Rick-
er 1973 for discussion). Non-transformed power equation for
relationship is W = 3.8 x 10~6 L3-'5, where W = weight in kg
and L = total length in cm.

al conditions was approximately 1 500 m. How-
ever, practical working distances were much less
due to transmission loss in the shallow waters
around the reef (20-30 m deep). Ranges were
estimated by relative audible strength calibrated
prior to tracking sessions.

A set of cursory experiments were performed
to test the sensitivity of sharks to weak electric
fields. Sharks attracted to the boat by chum were
presented two pieces of bait, approximately one
meter apart, suspended on the surface from lines
attached to the end of 7-m bamboo poles. Two
saltwater electrodes, similar to those used by
Kalmijn (1978), were attached to one bait (the
experimental). Electrodes consisted of one-meter
lengths of 6.3 mm inside diameter tygon plastic
tubing filled with a 3% seawater-agar gel. One
end of each tube was open to the water, while at
the other end a 32-mm stainless steel pin with
wire lead was inserted. Lead wires (+ and — )
were connected to a Grass S-6 stimulator. The
two saltwater electrodes were attached behind
the experimental bait and spaced 10 cm apart.
The control consisted of bait only. The experi-
mental bait was presented in two different ex-
periments that used 1) pulsed (2.2 volts at source,
5 Hz, 1.9 ms duration) current, and 2) constant
(0.5 and 2.2 volts at source, DC) current elec-
trical fields. Each test sequence began when a
shark visually oriented to and approached the
baits. Once the shark was within 2 m of the baits,
the stimulator was turned on and choice of bait
fed upon recorded. Relative positions of the con-

trol and experimental baits were randomly de-
termined to control for extrinsic cues.

RESULTS AND DISCUSSION

SIZE.— The length- weight relationships of 127
white sharks, based on records at the California
Academy of Sciences and those of published and
contributed sources, are shown in Fig. 3. The
largest shark record we found was that of a 6.4-
m-long (21 ft), 3324-kg (7302-lb) individual cap-
tured off Cojimar, Cuba, in 1945 (Guitart and
Milera 1974). The maximum size previously re-
ported for a white shark originated from an in-
correct record of an 1 1 . 1 -m individual from Port
Fairy, Australia, reported by Giinther (1870).
Randall (1973) re-examined the jaws of this spec-
imen and concluded that the correct total length
was approximately 5 m, well within the size dis-
tribution of sharks currently on record. The pur-
ported capture of a 9-m (29.5-ft) white shark said
to be from Vila Franca, Azores, is probably er-
roneous (see Ellis 1983). The smallest published
record was a 125-cm (49-in), 20-kg (44-lb) spec-
imen reported by Smith (195 1). Robert Johnson
(pers. comm.) has advised us of three juveniles
captured offBaja California that ranged from 1 30
to 135 cm (51 to 53 in) total length and weighed
less than 18.2 kg (40 Ib).

Remarkably, we found no well-documented
records of female white sharks with fetuses or
pups. Bigelow and Schroeder (1948) reported
embryos ranging in length from 20 to 61.6 cm,
but gave no further source information. One fe-
male taken near Alexandria, Egypt, was reported
to have nine embryos, each 0.61 m (2 ft) long
and weighing 49 kg (108 Ib). This erroneous
weight probably represents a total for all nine
embryos, and translates to a more reasonable
mean of 5.4 kg (12 Ib) for each fish. We can only
speculate about this lack of pregnant females in
the capture record. Females may pup in less fre-
quently sampled areas, such as remote geograph-
ic regions, oceanic waters, or deeper pelagic hab-
itats. Although it is possible that females fast
while pregnant, this would not completely ac-
count for the phenomenon, because many of the
largest females on record were taken by harpoon
rather than with bait and hook. Perhaps pregnant
females undergo spontaneous parturition when
hooked or harpooned and therefore eliminate key
embryonic evidence before they are landed. More
critical examination of the reproductive tract of

TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK

225

FIGURE 4. Still photographs of white sharks feeding on baits
near Dangerous Reef, Australia, (a) Shark begins to raise snout
and depress lower jaw. (b) Mouth opened fully with head and
snout raised, (c) Palatoquadrate protrusion and lower-jaw el-
evation, (d) Mouth closed; head is raised and disassociated
from upper jaw. (e) Head and snout drop to normal position.
Photos a, c, d, e by T. Tricas. Photo b by P. Romano.

freshly landed specimens might provide useful
insight to this enigma.

FEEDING ETHOLOGY.— The following section is
based on our observations and the analyses of
films taken of white sharks feeding on bait at
Dangerous Reef, Australia. Although baited sit-
uations can only simulate natural conditions, the
feeding behaviors observed in these sessions rep-

resented natural patterns because white sharks
normally take prey at the surface (Ainley et al.
1981; personal observations).

The following descriptions of the structures
and mechanics involved in biting actions of white
sharks employ terminology similar to that used
by Moss (1972, 1977). Cinemaphotographic
analysis of 36 feeding bouts revealed five basic

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14

components that constitute a single feeding ac-
tion.

1) Snout Lift: This movement involves an up-
ward lifting of the snout and head, and initiates
the feeding action (Figs. 4a and b, 5b). The degree
of snout lift ranged from a slight upward move-
ment to a pronounced elevation that produced
an acute angle behind the head (30-40° above
the longitudinal body axis). The intensity of snout
lift varied in relation to size of bait, angle of
approach to the bait, and possibly to level of
motivation (e.g., hunger).

2) Lower-Jaw Depression: Like the snout lift,
lower-jaw depression occurs at the start of a feed-
ing action. It is characterized by a ventro-pos-
terior movement of the tip of the lower jaw (Figs.
4a and b, 5b). This motion, along with the snout
lift, fully extends the gape.

3) Palatoquadrate Protrusion: Closure of the
mouth is marked by disassociation of the upper
jaw from its original juxtaposition ventral to the
cranium, and subsequent protrusion out of the
oral cavity. The upper jaw rotates in an antero-
ventral direction, while the snout remains at its
elevated position (Figs. 4c, 5c). During palato-
quadrate protrusion the teeth become fully ex-
posed and are directed downward. Eversion of
the upper jaw was readily visible by exposure of
the reddish connective tissue on the surface of
the jaw cartilage.

4) Lower-Jaw Elevation: Concurrent with the
initiation of palatoquadrate protrusion, the low-
er jaw begins an antero-dorsal (upward) motion
(Figs. 4c, 5c). These movements collectively pro-
duce the closing action of the jaws.

5) Snout Drop: After single-bite feeding bouts
the snout returns to its normal pre-feeding po-
sition. This results from a drop of the head and
snout, and a retraction of the palatoquadrate car-
tilage to its position immediately ventral to the
cranium (Figs. 4e, 5d). During multiple-bite bouts,

FIGURE 5. Components of a feeding action pattern. (A) Shark
just prior to initiation of feeding action. Snout and lower jaw
are at normal resting position. (B) Snout lift and lower-jaw
depression result in maximum gape. (C) Palatoquadrate pro-
trusion rotates upper jaw forward and downward exposing
upper teeth. Lower jaw moves forward and upward. These two
components comprise the actual bite. (D) Snout drop entails
retraction of palatoquadrate cartilage to its normal juxtapo-
sition beneath cranium. Snout drop occurs at the end of a
feeding bout and is not an essential component of the biting
action. Arrows indicate direction of jaw movements.

TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK

227

Maximum
Gape

I rB

i[E

1 rB

^ U

.5 .6

Time (s!

1.0

1.2

FIGURE 6. Timing of feeding actions for eleven consecutive bites made by a 3.5 m (TL) white shark. Mean times indicated
by dots. Horizontal lines show 95% confidence limits. Key: B = begin, D = depression, E = end, Elev = elevation, LJ = lower
jaw, PQ Prot = palatoquadrate (upper jaw) protrusion, S = snout.

the snout remains partially elevated prior to the
next biting action (Fig. 4d). The retention of an
elevated snout in these cases results in shorter
time intervals between bites.

Mean durations for components of eleven
complete successive feeding events recorded for
one shark are shown in Figs. 6 and 7. Depression
of the lower jaw was the fastest component (x =
0. 140 s), and the snout drop duration the longest
(x = 0.405 s). Total time for a complete biting
action, including the snout drop, ranged from
0.750 s to 1.708 s (x = 0.985 s). Temporal anal-
yses of film footage showed that the sequence of
each feeding component fell in a fixed order with
a non-overlapping range of time limits for each
individual head and jaw movement. While each
action showed a range in timing, minimal over-
lap was detected between events. The four com-
ponents occurred within a mean time of 0.443
s, and never was a shark observed to partially
complete a bite once the snout lift and lower-jaw
depression actions were initiated. The snout drop,
however, was not always an integral part of a

feeding action, except at the termination of a
feeding bout (as discussed above), and may be
subject to sensory feedback or motivational
changes.

Our observations on the mechanics of jaw pro-
trusion in the white shark are similar to those of

S Lift

L J Depress

LJ Elev

0)
Q.
D

S Drop

Time (s)

i.O

FIGURE 7. Range of durations for feeding events. Data from
same shark as in Fig. 6.

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14

Alexander (1967) on Squalus. He too found a
head lift component to precede jaw eversion. This
action is initiated by contraction of the muscles
at the posterior region of the head and may lead
to a pronounced snout lifting prior to feeding in
other sharks (Backus et al. 1956; Moss 1972).
Actual jaw protrusion in the white shark begins
after maximum gape is achieved and the mouth
begins to close (Figs. 4, 5). In fact, full protrusion
of the palatoquadrate cartilage is not achieved
until about midway through the jaw closing ac-
tion. Once the mouth begins to close, palato-
quadrate extension is very fast (x = 0.083 s, n =
1 1) and represents the actual downward move-
ment of the fully exposed teeth during a bite.

The mechanics and function of the protrusible
jaw in large sharks has played a major role in the
evolution of their feeding habits (see Moss 1977
for review). Special hydrodynamic problems ex-
ist for non-demersal sharks because of their lack
of a gas-filled swim bladder (but see Bone and
Roberts 1969) and maneuverable paired fins
(Alexander 1967). The general streamlined body
form is considered an evolutionary response to
this problem (Alexander 1967; Budker 1971;
Thomson and Simanek 1977). The development
of a protractile jaw has allowed large lamnid and
carcharhinid sharks to retain a hydrodynami-
cally efficient fusiform body and the capacity to
take clean bites with a subterminal mouth. The
rounded pattern of bites taken from prey too
large to swallow whole comes primarily from the
upward and forward rotation of the lower jaw
that secures the mouth to the prey, and the down-
ward and forward cutting rotation of the upper
jaw. The detached hyostylic association of the
upper jaw and chondrocranium also permits the
upper jaw to close downward much faster than
it could if it had to pull the head with it as it
closed. This rapid downward movement of the
massive unattached upper jaw produces a strong
resultant force that facilitates the cutting action
of the serrated teeth.

PREDATORY BEHAVIOR. — Sharks used various
capture modes to take baits depending on the
bait's size and its position relative to the surface.
In situations where large pieces of meat were
suspended or floating at the surface, two com-
mon approaches were observed.

1) Underwater Approach: In this behavior,
sharks swam parallel to and approximately 0.5
m below the surface until less than 1 m away from

the bait. In situations where bait was freely float-
ing on the surface, sharks swam at normal swim-
ming speeds as the prey was engulfed. In cases
where the bait was suspended by pole and line,
sharks would typically bite the bait and attempt
to pull it under by depressing their heads. Sharks
that did not sever the line would often hang ver-
tically and repeatedly bite at the bait, displaying
all components of the bite behavior. Sharks were
persistent in attempts to take the bait after an
attack was made.

2) Surface Charge: The second, less common
feeding behavior on bait at or near the surface
was a rapid accelerated rush. Here a shark would
approach and engulf the bait as it swam by at a
relatively fast rate. This behavior was most com-
monly observed on newly arrived sharks in an
excited state. Unlike the underwater approach,
a charge was made at the surface. This behavior
created considerable disturbance well before the
bait was taken. Charging behavior of a similar
nature was described for the blue shark (Prionace
glaucd) feeding on dense surface schools of squid
(Tricas 1979).

White sharks were also observed feeding un-
derwater, and exhibited different predatory be-
haviors than when taking prey from the surface.
Two additional modes are presented.

3) Normal Underwater Pass: This feeding be-
havior was observed when a shark approached
a relatively small submerged bait. Sharks ap-
proached with the mouth opened wider than dur-
ing normal swimming and raised the snout slightly
when approximately 1 m from the bait. When
the bait contacted the underside of the snout, the
lower jaw depressed slightly and the bait was
taken. In this behavior the snout lift and lower
jaw movements were present, but not as pro-
nounced as in surface feeding modes, and there
was no protrusion of the upper jaw. The under-
water pass appeared to be first mediated by vi-
sion prior to contact, and second by tactile sen-
sory input when the snout touched the prey just
prior to initiating a feeding action. Additional
sensory systems (e.g., gustatory or electrorecep-
tive) may also be involved in normal feeding
situations at close ranges.

4) Side-Roll: A similar approach to a normal
underwater pass occurred where a shark rolled
onto its side just prior to engulfing the submerged
prey. Here the shark maintained its horizontal
approach until approximately 1-2 m away from

TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK

229

the bait. It then rolled aproximately 60° from
normal, took the prey, and returned to an upright
swimming attitude.

These latter two approaches involved no de-
tectable change in swimming speed and em-
ployed the characteristic movement sequence of
head and mouthparts.

Numerous observations on the variability in
feeding patterns of sharks in relation to prey type
and feeding conditions exist. Budker (1971) re-
ported that in normal feeding situations sharks
exhibit no body contortions when they consume
small fish prey which are swimming at their own
level or slightly below. This appears to be the
case for white sharks taking small pieces of bait
in normal underwater passes. He further states
that there are only two situations that might re-
quire a different type of approach to a bait. These
are either when a bait is attached to a hook and
the shark must turn on its side to avoid the line
with its snout, or when bait is floating and the
shark must thrust its snout out of the water to
get its mouth around the bait. We agree with his
conclusion in regards to floating baits, with the
addition that this includes natural feeding situ-
ations as well. This behavior has been observed
for white sharks feeding on pinnipeds at the sur-
face (Ainley et al. 1981; personal observations),
and for tiger sharks (Galeocerdo cuvief) feeding
on both surface baits and normal prey (see Moss
1972; Gilbert 1963 for pictures). In addition, it
is clear that side-roll behaviors may also occur
in natural feeding situations, and are not nec-
essarily responses to obstructions during feeding.
Tricas ( 1 979) found that blue sharks approached
small, moving anchovy baits from behind and
took them in a normal swimming posture, while
larger whole mackerel baits were taken from be-
hind by sharks that partially rolled onto their
sides. This variation was attributed to the size
of the prey and its position relative to the mouth
just prior to capture.

Observations of white shark feeding behavior
are limited to artificial situations in which sharks
were attracted to feed on tethered fish or horse-
meat (this study) and the few instances when
white sharks were observed feeding on dead ce-
taceans (e.g., Pratt et al. 1982). Some information
has been gained from interviews with shark at-
tack victims, although most of these did not see
the shark before or during the attack (cf. Miller
and Collier 1981) and may have made biased

observations. White sharks have been kept alive
for short periods in large aquariums, but none
have attempted to feed (McCosker 1981).

On the basis of information discerned from
white shark attacks on pinnipeds and humans,
and our observations of their feeding on bait, we
can best summarize the predatory attack strategy
as follows. An adult white shark is not agile
enough to capture a fleeing, darting pinniped;
hence, it generally attacks its prey by surprise.
Bite scars on northern elephant seals (Mirounga
angustirostris), California sea lions (Zalophus
californianus), Australian fur seals (Arctocepha-
lus doriferus) (Fig. 8), and sea otters (Enhydra
lutris) (Fig. 9, also see Ames and Morejohn 1 980)
are usually located on the ventral region of the
body (e.g., haunches and flippers). This indicates
that attacks were made from behind and beneath
the prey. A typical attack scenario might entail
a shark swimming a few meters beneath the sur-
face, searching for the silhouette of a pinniped
or sea otter at the surface. Once a prey is sighted,
the shark ascends and at close range (approxi-
mately 1 m) begins a feeding action as described
above. After attacking large prey such as an el-
ephant seal, the shark probably routinely retreats
a short distance from the injured (and at least
partially immobilized) prey and swims cautious-
ly within the area, apparently waiting for the pin-
niped to bleed to death or lapse into shock. Dur-
ing the attack, white sharks often roll their eyes
posteriorly, which reduces the risk of injury to
the eye by the teeth or nails of a struggling prey.
The retreat behavior is also adaptive since it
eliminates the chance of injury via contact after
the initial attack is made. This "bite and spit"
strategy might explain why seals that have es-
caped after attack usually have a single massive
bite. This might also provide insight into the
paradox of why humans are rarely consumed
after being attacked. Since humans rarely dive
or swim alone, the victim is usually quickly res-
cued or removed from the attack area by others,
precluding a second attack. The white shark-
related human fatalities that have occurred in
California and Oregon waters since 1926 (5 of
40 attack victims) have all resulted from trau-
matic blood loss and did not involve massive
consumption by the shark. This alternative the-
ory might be more reasonable than the sugges-
tion that humans are "distasteful" to white sharks,

230

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14

«r>

"15^
% <&*-<

FIGURE 8. Wounds inflicted upon pinnipeds by white sharks. It is possible to hypothesize the posture of prey and the attack
behavior of the shark from bite scars. Elephant seals (Mirounga angustirostris): (A) Subadult survivor at Afio Nuevo Island,
California. Note lower jaw puncture wounds and tearing caused by upper jaw teeth. Photo by R. Bandar. (B) Adult female
survivor at Southeast Farallon Island, California. Again note lower and upper teeth wounds. Photo by S. H. Morrell. California
sea lions (Zalophus californianus): (C) Adult survivor at Afio Nuevo Island. Photo by R. Bandar. (D) Subadult male carcass
(left) and Richard Ellis (right) at Afio Nuevo Island. Photo by Pam Wing. (E) Subadult male carcass along the central California
coast. Photo by R. Bandar. Southern fur seal (Arctocephalus doriferus): (F) Large adult male survivor at South Neptune Island,
South Australia. Photo by J. McCosker.

particularly when one considers the euryphagic
diet of the fish.

PREDATOR-PREY RELATIONSHIPS.— The stom-
ach contents of nine white sharks (193-51 1 cm
total length) captured in northern and central
California waters are presented in Fig. 10. Seven-

ty-eight percent of the sharks had recognizable
food items in their stomachs. The most frequent
prey was the California bat ray (Myliobatis cali-
fornica), found in four stomachs; other fish prey
were less frequent in the diet. Fifty-six percent
of the sharks examined contained elasmo-

TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK

231

FIGURE 9. Sea otters (Enhydra lutris) from the central California coast. Above, adult in normal feeding or basking posture
along the edge of a kelp bed in Monterey Bay. Photo by J. McCosker. Below, lacerated carcass from which several white shark
tooth fragments were removed, suggesting that the animal was bitten at the surface while in a belly up, prone position. Pismo
Beach. Photo by J. Ames.

branchs, and 44 percent contained teleost prey
species. No evidence of predation on marine
mammals was found in the nine sharks.

Although the white sharks took prey that nor-
mally occur in both pelagic and inshore habitats,
the two most frequent prey are generally asso-
ciated with demersal inshore communities. The
California bat ray (M. californicd) is common in
bays and inshore sandy habitats 2-50 m deep,

where it feeds on benthic sand-dwelling inver-
tebrates. The spiny dogfish (Squalus acanthias)
is also demersal, being found in both shallows
and deeper offshore waters. Other prey species
that live on the bottom in inshore areas are the
lingcod (Ophiodon elongatus) and the cabezon
(Scorpaenichthys marmoratus). These latter two
species are relatively sedentary, have small home
ranges, and show cryptic coloration. Limbaugh

232

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14

Myliobatis California IE)
Squ.lu. acanthi, 3 IE)
Cetorhinus maxlmut IE)
Cynoscion noblll*
Galeorhintis zyopterus (E)
Ophlodon elongatus

% Occurrence

FIGURE 10. Stomach contents of nine sharks captured in
northern California waters on record at California Academy
of Sciences. % occurrence = percentage of the nine shark stom-
achs that contained that prey item. E = elasmobranch, all oth-
ers are teleosts.

(1963) reported cabezon from the stomachs of
three immature sharks captured at La Jolla, Cal-
ifornia, and described a number of incidents that
indicate 5". marmorafws is an important prey for
young white sharks. Earlier researchers ques-
tioned how sharks could detect and capture such
inconspicuous and apparently inaccessible prey;
our studies suggest that weak electric fields might
be involved in prey detection (see Sensory Bi-
ology section below).

The white sea bass (/l/rac/osr/on (=Cynosciori)
nobilis) also occurs in shallow rocky inshore hab-
itats, and is often found among canopies of giant
kelp (Macrocystis pyrifera). Unlike the majority
of the other inshore prey species, however, it
occasionally swims in the water column as well
as on the bottom.

White sharks have been reported to feed on
the carcasses of captured basking sharks (Fast
1955), although we know of no published ac-
counts of predation under natural circumstances.
However, potential vulnerability of basking
sharks to large predators was suggested by Lim-
baugh (1963) in an account of a dead basking
shark with a large wound probably inflicted by
killer whales. Basking sharks, which reach lengths
of more than 1 1 m, are found seasonally in off-
shore waters of central and northern California.
From aerial surveys made over a 2.5-yr period
near Monterey, California, Squire (1967) found
that basking sharks were most common from
September through May, when water tempera-
tures were generally below 14°C. White shark

Fish

Pinnipeds
Cetaceans
Other prey

LJ Elasmobranchs

LJ Actinopterygians

% Occurrence

FIGURE 11. Stomach contents of 33 white sharks. Data
combined from this study and other published records. % oc-
currence = percentage of the 33 sharks that contained the prey
category. Fish prey subdivided into elasmobranchs and rayed-
fin fishes (teleosts and sturgeons). Other prey include birds,
crustaceans, and sea turtles.

sightings, however, were most common in the
warmer-water months of May through August,
when water temperatures neared or exceeded
1 4°C. The cause of the seasonal disappearance
of basking sharks from the coastal waters of Cal-
ifornia remains unknown. Other prey that in-
habit pelagic waters include the soupfin shark
(Galeorhinus zyopterus), the Pacific sardine (Sar-
dinops sagax), and occasionally bat rays (Myl-
iobatis californicd) (Roedel and Ripley 1950;
Federetal. 1974).

Combined data on the food habits of 33 white
sharks from this study and other published rec-
ords are shown in Fig. 1 1 . Here again, fish were
the most frequent prey items, occurring in over
half of white sharks in the analysis. Elasmo-
branchs and rayed-fin fishes (teleosts and stur-
geons) comprised equal proportions (each oc-
curred in 30 percent of sharks analyzed) of the
piscine prey. Pinnipeds were also a major com-
ponent in the diet of sharks, while cetaceans and
other prey groups were less common. Bass et al.
(1975) provided the only other gut content data
from white sharks useful for comparison. They
too found both elasmobranchs (40 percent of
sharks examined) and teleost fishes (25 percent)
as the most common prey items, although little
information was given on specific identification.

Figure 1 2 shows the distribution of fish and
mammal prey in relation to shark size. Fish prey
predominated in the diet of sharks approxi-
mately 3 m or less (TL), while pinnipeds and
cetaceans predominated in those of larger sharks.
This shift in diet may occur for a number of
reasons. For example, larger sharks are less agile
and would be less successful in chasing and cap-

TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK

233

O Pinnipeds & Cetaceans
• Fish

o oooooo oo o o

100 200 300 400 500 600
Total Length (cm)

FIGURE 12. The relationship between white shark length
and prey type. Data taken from stomach contents of the 33
specimens in Fig. 1 1 .

turing smaller fish prey that dart about when
pursued. Larger sharks may thus switch to dif-
ferent prey types and associated new hunting
modes. In addition, the energetic requirements
of large, warm-bodied sharks may be better met
by prey high in fat content (i.e., high-energy-
density prey). Carey et al. (1982) estimated the
metabolic rate for a 4.6-m white shark, and con-
cluded that the animal could survive for ap-
proximately 1.5 months on 30 kg of whale blub-
ber (a conservative meal size). They suggest this
to be adaptive during long intervals between en-
counters with prey. Although little is known of
the movements of white sharks, they do show
seasonal peaks in abundance in California waters
(Squire 1967; Ainley et al. 1981), which might
indicate some sort of regional or long-distance
movement.

Morphological differences between large and
small sharks may also account for different pred-
atory tactics. Fig. 1 3 shows the relationship be-
tween tooth shape and shark total length. Smaller
sharks have a relatively long, narrow tooth shape
that is better adapted for grasping prey like small
fishes. This feature is so well developed in small
white sharks that they are often incorrectly iden-
tified as mako sharks (hums spp.) (Smith 1951,
1957). At about 3 m TL, the teeth broaden at
the base and take on the diagnostic triangular
serrated form. Unlike the long narrow teeth, this
shape is well-suited for gouging and cutting pieces
from prey too large to swallow whole. Le Boeuf
et al. (1982) found evidence that marine mam-
mals were the only prey of large white sharks
they examined from California. Of seven spec-
imens examined, all but one were approximately
4 m or longer and had evidence of marine mam-
mals in their stomachs. The only exception was
the smallest shark (2.4 m TL), which had only a
10-cm patch of pinniped pelage in its stomach.

Total Length ( m)

FIGURE 1 3. The relationship between shark total length and
tooth shape. Tooth shape expressed as the ratio of width of
enamel base to medial height of enamel for the first tooth, right
side, upper jaw of 16 sharks. Low ratio indicates a long narrow
tooth shape; higher ratio indicates relatively broad triangular
shape.

Perhaps this shark's teeth were too narrow to
excise a portion of flesh.

In California waters, elephant seal populations
at offshore rookeries peak in both the spring and
winter months (Le Boeuf et al. 1 974), but almost
no predation occurs during the spring peak. Hy-
pothetical explanations advanced to explain this
seasonal discrepancy in predation include either:
1 ) sharks fasting while breeding; 2) water too cold
for sharks to feed; or 3) emigrations of sharks
from the area. Even though sharks occur in Cal-
ifornia waters during the spring (Miller and Col-
lier 1981), the decrease in shark attacks is prob-
ably due to emigrations of large sharks from
coastal areas (see Squire 1967). Adult male seals
are more susceptible to shark predation because
they spend more time in the water near the rook-
ery during the breeding season than do females
(Le Boeuf et al. 1982). It is possible that the loss
of peripheral males to sharks may not adversely
affect the population because of the polygynous
mating system of the elephant seal, where rela-
tively few dominant males do the majority of the
breeding.

Although it is clear that white sharks do nor-

234

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14

FIGURE 14. Underwater photo of a male white shark (ap-
proximately 3.5 m TL) in a "tail stand" posture with snout
directly over zinc anode on rudder of study vessel. Photo by
T. Tricas.

mally prey upon elephant seals, the significance
of the interaction is not evident. Ainley et al.
(1981) reported an increase in the number of
attacks on elephant seals at the Farallon Islands
between 1970 and 1979, but their data indicate
a density-dependent relationship between num-
ber of attacks and numbers of elephant seals.
More data are needed on the mortality rates of
attacked seals and on numbers in the shark pop-
ulation before any effects of shark predation on
elephant seal populations can be quantitatively
assessed.

SENSORY BIOLOGY.— Our cursory field exper-
iments and observations qualitatively indicate
white sharks are sensitive to electric fields. In the
pulsed electric field tests, sharks took the exper-
imental bait 8 times (73 percent) and the control
3 times (27 percent). In the constant current (DC)
tests the experimental was taken 4 times (44 per-
cent) and the control 5 times (56 percent). Al-
though our sample size was too small to show
any statistically significant preference for baits
with either type of electric field, sharks did take
baits with the pulsed electric field almost three
times more often than the control. The sharks
also appeared to be more responsive to pulsed
fields than to continuous fields. Kalmijn (1971,
1974) reported that sharks were most responsive
to weak electrical fields at frequencies from 0
(DC) to 8 Hz.

We also observed the behavior of sharks to
metallic objects attached to the bottom of the
boat. On three occasions one of us (TCT) watched

from underwater a 3.5-m shark approach zinc
plates attached to the boat's rudder and assume
a near vertical "tail stand" posture (Fig. 1 4). The
shark remained upright for approximately 10-
20 s as it waved its snout approximately 5-10
cm above the zinc. Sharks were also observed
several times to swim back and forth with their
snouts very near a 1 0-m-long copper grounding
strip on the bottom of the boat's hull.

We interpret these observations as a response
by sharks to the galvanic currents produced by
the electrochemical interaction between the me-
tallic plates and seawater. White sharks have a
well-developed system of ampullae of Lorenzini
(Fig. 15), and although the role of electric detec-
tion of prey by sharks is well demonstrated (see
Kalmijn 1978, 1982), the degree of importance
for such a sensory modality in white sharks re-
mains unknown. It is noteworthy, however, that
electric fields produced by large mammals (e.g.,
humans and presumably pinnipeds) in seawater
are well within the sensory range of elasmo-
branchs (Kalmijn 1971). Perhaps young white
sharks are able to detect electrically sedentary
camouflaged fish prey like the cabezon (Scor-
paenichthys marmoratus). It also seems reason-
able that the ampullae would be particularly use-
ful to detect: 1) the location of a marine mammal
at the moment just prior to attack; 2) any change
in position or escape attempts by the prey; and
3) any change in the prey's condition, such as
bleeding, which might alter the strength or sig-
nature of the electric field.

TELEMETRY.— Two sharks were tagged with
temperature-sensing transmitters during this
study. The first shark (a 4.5-m male) carried a
unit that monitored ambient water temperature
only. After tagging, the shark remained around
the boat even after all baits were removed from
the water. The boat was then moved away from
the area and the shark began to move westward;
parallel to the north shore of Dangerous Reef.
Once past the island the shark moved offshore
in a northwesterly direction. Contact was lost
with the animal approximately 4 h after initial
tagging, due to its rapid speed and bad seas that
created poor tracking conditions. During this time
the shark swam in waters 20-2 1°C as indicated
by the temperature sensor on the transmitter.

The second shark was tagged on 22 January
1980. The body temperature probe was placed
31 cm deep into the lateral musculature, ap-

TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK

235

FIGURE 15.
C. J. Slager.

Distribution of the ampullae of Lorenzini on the head of a young female white shark (CAS 37917). Figure by

proximately 25 cm below the first dorsal fin. This
shark was monitored continuously near the boat
for approximately 2 hr, until it swam out of range.
It returned to the anchored boat near midnight,
and then again departed. Results of the thermal
data are presented in Table 1 and Fig. 16. The
shark swam in water ranging from 20.9° to 2 1 .5°C.
Mean difference between ambient and body tem-
perature was 3.7°C, and ranged from 3.2° to 4.3°C.

TABLE 1. EPAXIAL MUSCLE TEMPERATURES OF A 3.5 M (TL)
MALE WHITE SHARK MONITORED AT DANGEROUS REEF, SOUTH
AUSTRALIA ON 22 JANUARY 1980. Mean (AT) = 3.7°C. SD =
0.37.

Measure-
ment

Temp (°C)

Water

Body

Difference
(AT)

21.2

25.2

4.0

21.5

24.7

3.2

21.2

24.7

3.5

21.2

25.2

4.0

21.2

25.2

4.0

21.2

25.2

4.0

21.2

24.7

3.5

20.9

25.2

4.3

20.9

24.7

3.8

20.9

24.2

3.3

20.9

24.2

3.3

20.9

24.2

3.3

Largest and smallest differences were recorded
when the shark entered water of a different tem-
perature, before internal temperatures could con-
form. This time lag to thermal equilibrium and
variation in muscle temperature indicate that the
shark did not thermoregulate. Carey et al. (1982)
found that a 4.6-m white shark had a body tem-
perature 3-5°C higher than the surrounding water.
Their shark swam over deeper waters, and for
the most part remained in the thermocline. Tem-
peratures were lower in their study, ranging ap-
proximately from 5° to 19°C ambient, and 18°

WHITE SHARK MUSCLE TEMPERATURE

White Shark Epaxial Musculalu

FIGURE 1 6. Temperature difference between ambient sea-
water and epaxial musculature of a 3.5 m TL white shark,
monitored on 22 January 1 980 at Dangerous Reef, South Aus-
tralia. Question marks (?) indicate time interval when shark
swam away from anchored study vessel and out of telemetry
range. Figure by K. O'Farrell.

236

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 14

FIGURE 17. (Right) Silhouette of a surfer on a contemporary surfboard. (Left) Silhouette of an adult female (TL = 1.7 m)
harbor seal (Phoca vitulina). Photo by Al Giddings.

to 23°C muscle temperature. Our study took place
in relatively shallow waters around Dangerous
Reef (< 30 m), and we found no sign of a marked
thermocline. The water and shark-muscle tem-
peratures we recorded were generally higher
(20.9°-2 1.2° and 24.2°-25.2°C, respectively), but
they are consistent with the values for body tem-
perature elevation over ambient recorded by Ca-
rey et al.

One of the primary advantages of being warm-
bodied is thought to be related to the changes in
muscle physiology as temperature increases. It is
known that a 10°C increase in temperature may
result in a three-fold increase in the contraction-
relaxation rate of frog muscle (Hartree and Hill
1921). For fish, this may be translated to an in-
crease in potential tail-beat frequency and a re-
lated increase in sustained swimming speed.
Higher speeds may be selectively advantageous
when chasing prey or fleeing from predators. In
addition, conservation of heat theoretically al-
lows for more total energy conversion to work,
thus enabling an animal to swim longer distances
on a given meal. Being warm-bodied might also

allow for temporary excursions into colder or
deeper waters. This thermal inertia (see Neill et
al. 1976) would not only expand the range of
environments which the animal could exploit,
but would also permit increased swimming ef-
ficiency for predation at otherwise limiting en-
vironmental temperatures.

ON WHITE SHARKS AND SURFBOARDS. — In con-
clusion, we comment on the increasing attacks
by white sharks upon humans who surf in the
north Pacific. Since 1972, there have been 11
recorded white shark attacks upon surfers in Cal-
ifornia and Oregon (Miller and Collier 1981) and
one such attack in Hawaii in 1959 (Balazs and
Kam 1981). The similarity in appearance of the
silhouette of a prone human on a surfboard or
"belly board" to a large surface-basking pinniped
is clear (Fig. 1 7), and observations of attacks by
sharks upon surfers fit well with our assessment
of the feeding strategy of white sharks. Attacks
have occurred in the vicinity of pinniped rook-
eries, such as the much-publicized death of Lewis
Boren on 19 December 1981 at Spanish Bay,
Monterey, Calfornia.

TRICAS AND McCOSKER: PREDATORY BEHAVIOR OF THE WHITE SHARK

237

Since the early 1970s, the trend in surfboard
design has been toward an increase in flotation,
reduction in board length, multiple posterior-
fixed rudders ("skegs"), and bifurcated or "V"
tails. All of these modifications have enhanced
the similarity between the silhouette of a surfer
and that of a pinniped, and we suggest that this
may increase the probability of attack of surfers
encountered by white sharks. We feel it advisable
that those who surf be aware of and consider the
potential risks of surfing in coastal waters known
to be frequented by white sharks.

ACKNOWLEDGMENTS

We are particularly grateful to Al Giddings,
President of Ocean Images, Ltd., for financial
assistance, for providing access to his film library
for cinematographic analyses, and for sharing his
observations of shark behavior with us. We also
thank Terry Thompson, Ocean Images, Ltd., for
his assistance. Additional shark data were pro-
vided by W. I. Follett (CAS), L. J. V. Compagno
(CAS), J. Randall (Bernice P. Bishop Museum),
and the staff of the Department of Ichthyology
of the CAS. A. Dizon (NMFS, Honolulu) and I.
Cooke (Bekesy Laboratory, U. of Hawaii) pro-
vided facilities during construction of our trans-
mitters. We thank H. Tricas, C. J. Slager, K.
O'Farrell, S. Middleton, and S. Nakamura for
assistance with our figures; J. Ames, R. Bandar,
R. Dunne, A. Giddings, S. Morrell, P. Romano,
and P. Wing for allowing us to use their pictures;
and Bob Britcher and Chico Chingwidden, the
Master and the mate of the Nenad. We give spe-
cial thanks to Rodney Fox, for his guidance in
the field in South Australia and for helpful dis-
cussions concerning shark behavior, and to
Leighton Taylor, Jr. (Waikiki Aquarium), and
Phil Motta (Univ. of Montana) for their critical
reading of this manuscript.

Senior authorship of this paper was deter-
mined by the outcome of a pinball match played
at Port Lincoln, South Australia, in January 1 980.

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THOMSON, K. S., AND D. E. SIMANEK. 1977. Body form and
locomotion in sharks. Amer. Zool. 17:343-354.

TRICAS, T. C. 1979. Relationships of the blue shark, Prionace
glauca, and its prey species near Santa Catalina Island, Cal-
ifornia. Fish. Bull. 77:175-182.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 15, pp. 239-248, 5 figs., 1 table

September 19, 1984

DENDRODOA (STYELOPSIS) ABBOTTI, SP. NOV.

(STYELIDAE, ASCIDIACEA) FROM THE PACIFIC COAST OF

THE UNITED STATES, AND ITS IMPACT ON SOME GONADAL

CRITERIA OF ITS GENUS AND SUBGENUS

Andrew I odd Newberry
Cowell College, University of California, Santa Cruz, California 95064

ABSTRACT: Dendrodoa (Styelopsis) abbotti, a newly described styelid ascidian from the central and northern
California coast, the San Juan Islands of Washington, and southwestern Vancouver Island, resembles I).
cornea but differs in branchial and gonadal traits. Inclusion of D. abbotti in the genus Dendrodoa requires
modification of the gonadal criteria of the genus to accommodate styelan gonadal resemblances (non-encap-
sulation of the testis-lobes with the ovary) and styelan or cnemidocarpan spermiducal resemblances (gonad's
single vas deferens and spermipore).

INTRODUCTION

The tunicate named and described in this pa-
per, Dendrodoa (Styelopsis) abbotti, is a styelid
ascidian that has long been collected along the
central and northern California coast. Donald P.
Abbott, who first found this ascidian in 1948
near Point Arena (Mendocino County), included
it as "Alloeocarpa sp." in the urochordate key of
the second edition of Light's Manual (Light et
al. 1954) but, for want of more certain identifi-
cation, omitted it from that handbook's third
edition (Smith and Carlton 1975).

This ascidian's aggregative habit does create
an appearance of budding (Fig. 1 A), but adjacent
zooids' tests are unfused and easily separated
from one another; no evidence of budding has
been found in several hundred zooids from sev-
eral sites and all seasons. Apparently, then, this
is a solitary ascidian and cannot be placed in the
genus Alloeocarpa. It shows Dendrodod's restric-

tion of the single, elongate ovary to the zooid's
right side. The ovary's unbranched shape and
the pharynx's simplicity place the species in the
subgenus Styelopsis of Dendrodoa. The specific
name, abbotti, honors Professor Donald P. Ab-
bott, of the Hopkins Marine Station of Stanford
University, who has shared with his students and
colleagues a singular keenness of intellect and
generosity of spirit, and it expresses the esteem
and affection of his fellow ascidiologists.

MATERIALS AND METHODS

This report is based principally on specimens
collected intertidally at Pigeon Point, San Mateo
County, California (lat. 37°11'0"N, long.
122°23'10"W), at intervals of roughly six weeks
throughout 1977. 1 have also drawn on material
taken over the past two decades from there; from
Point Pinos, Monterey County, California (lat.

[239]

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 15

FIGURE 1 . A) Living zooids of Dendrodoa abbotti, including one (upper left-center) that has been wounded or severely
disturbed and is extremely contracted while the others remain relaxed. Zooids are about 1 cm long. B) Closer view of two slightly
disturbed zooids, showing siphons in the process of bilabial closure. Brood pouch is detectable in the left zooid by inflated
aspect of the posterior region of the zooid (to left in photo). Zooids are about 1 cm long.

NEWBERRY: DENDRODOA (STYELOPSIS) ABBOTTI, SP. NOV.

241

36°38'0"N, long. 121°56'0"W); and subtidally
from Peavine Pass (lat. 48°35'4"N, long.
122°45'48"W) in the San Juan Islands, Wash-
ington. I have used, as well, D. P. Abbott's un-
published notes and drawings of specimens from
northern and central California.

In all, I have examined approximately 30 spec-
imens thoroughly. I have examined several doz-
en more in a cursory way to verify the criteria
that characterize the species. All specimens were
relaxed with MgCl2 or MgSO4, and menthol, then
fixed in seawater Bouin's fluid or 10% formalin,
and all were preserved in 70% ethanol. The
Bouin's-fixed material provided excellent serial
sections but brittle dissections. Formalin always
'fixed adequately for dissections but rarely well
enough for close scrutiny by serial section (which
was required, for example, to trace the very fine
spermiducts). Specimens were dissected in 70%
ethanol. Dissected specimens usually were
stained, once opened, with Grenacher's borax
carmine; serially sectioned specimens were either
prestained, often for prior dissection, in Gren-
acher's alcoholic borax carmine or stained in sec-
tion with "standard alum hematoxylin" (Galigh-
er and Kozloff 1964) and eosin. Prestaining
proved satisfactory for general examination, but
staining in section was necessary to reveal finer
structural details or to take advantage of the bet-
ter fixation achieved with Bouin's fluid than with
formalin.

COORDINATES.— The endostyle designates the
anterior-posterior axis and the ventral midline.
Thus, the dorsal midline extends from the- oral
siphon through and beyond the atrial siphon. By
these coordinates, the ovary lies against the right
ventral margin of the zooid, and the loop of the
gut dominates the left posterior region of the
zooid (Fig. 2).

DESCRIPTION OF SPECIES
Dendrodoa (Styelopsis) ahbotti, sp. nov.

TYPE-SPECIMENS. — Holotype at California Academy of Sci-
ences, San Francisco, Calif. (CAS #034790). Paratypes at Cal-
ifornia Academy of Sciences, San Francisco, Calif. (CAS
#034791).

TYPE-LOCALITY.— North side of Pigeon Point, San Mateo
County, California (lat. 37°1 1'0'N, long. 122°23'10"W).

OTHER RECORDS. — Intertidal records from Point Pinos and
Hopkins Marine Reserve (Monterey County), Pigeon Point
and Moss Beach (San Mateo County), Point Arena (Mendocino
County), California, and near Sooke, Vancouver Island, British
Columbia; subtidal records from Peavine Pass (San Juan Coun-
ty), Washington.

EXTERNAL APPEARANCE (Fig. 1 ). — Zooids round
or oval (lengthened antero-posteriorly) low
hemispheres; entire sub-endostylar surface ap-
plied to substrate; attached surface extends be-
yond ovary on right and gut-loop on left. Spec-
imens including test reach 8 to 1 2 mm length, 6
to 10 mm width, 2 or 3 mm height when relaxed;
zooids removed from test reach 8 to 10 mm
length, 6 to 8 mm width, 2 to 3 mm height. Test
clean, thin, and parchment-like, spreading as a
thin apron 1 to 2 mm wide on the substrate
around the zooid. Ventral test extremely thin.
Color in life translucent gray tinted with ochre
or very pale brownish pink, with borders of si-
phonal apertures sometimes slightly darker.
Zooids fixed in formalin become plain translu-
cent white-gray. Alive or fixed, zooid's branchial
sac, gut, ovary, and mass of brooded young are
faintly visible through dorsal and lateral regions
of test. Oral siphon far anterior; atrial siphon
placed centrally atop hemispheric zooid; both
siphons fairly evident in relaxed living animals
but reduced to obscure slits in contracted ones.
Relaxed zooids have circular siphonal apertures;
disturbed zooids close their siphons bilabially
into transverse slits (Fig. IB) and flatten them-
selves against the substrate within a delicately
crumpled test. Zooids are simple and non-bud-
ding but often aggregate in pairs or trios (rarely
groups of more) with young ones often settling
adjacent to or even on the test "apron" around
older zooids (but not on zooidal surfaces them-
selves). Mature zooids, even when tightly adja-
cent to one another, attach entirely to the sub-
strate itself; they do not form clumps of zooids
growing thickly one upon another. Adjacent
zooids often are oriented similarly on the sub-
strate.

VASCULAR ELEMENTS OF THE TEST.— Test- ves-
sels not prominent; as revealed by staining,
branching systems of test- vessels ramify toward
the margin of the test. Test- vessel ramifications
connect to zooid by one or more sub-zooidal
circulatory junctions; tips of all branches of test-
vessel ramifications end peripherally in slender,
bulbous vascular ampullae.

MANTLE.— Thin, lightly muscled mantle ex-
cept for extensive arrays of fibers radiating from
each siphon and controlling its bilabial closure;
fairly conspicuous concentric musculature sur-
rounding oral siphon, less developed concentric
musculature around atrial siphon. About a dozen
endocarps project from the mantle into the atrium

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 15

FIGURE 2. Dorsal view of zooid, with pharynx removed to show disposition of ovary and testis-lobes (including far posterior
group), vasa efferentia and vas deferens (including tiny spermiporal papilla near atrial siphon), mid- ventral endostyle and heart
beneath it, and gut-tract (stomach, pyloric duct, caecum, intestine). Position of oral and atrial siphons indicated by ovals.

FIGURE 3. Zooid opened by mid-ventral cut to show ovary, testis-lobes (including posterior lobes), larvae in brood pouch,
several endocarps (stippled), gut-tract, siphons with neural complex between them. Pharynx removed.

of most zooids; particularly large endocarps usu-
ally protrude from the atrial mantle anterior to
the ovary and in the region of the gut-loop.

ORAL TENTACLES. — 36 to 40 filiform oral ten-
tacles of three sizes; largest ones most abundant
(24-30), others about half their size intercalated
irregularly, a few to many tiny papillae evident
upon close examination of the band of oral ten-
tacles. Just distal to this circle of tentacles is a
siphonal flange that marks the inward limit of
the test that lines the oral siphon.

ATRIAL TENTACLES.— 40 to 50 tiny filiform
atrial tentacles in band analogous to that of the
circle of oral tentacles. Just distal to this incon-
spicuous circle is the atrial siphonal flange that
marks the inward limit of the test lining the atrial
siphon.

DORSAL TUBERCLE (Fig. 4).— Simple C-shaped
slit atop a short, stout projection; concavity of
the C faces posteriorly (toward the dorsal lami-
na). The dorsal tubercle is set slightly to the right
of the dorsal midline.

NEURAL COMPLEX.— In dorsal or ventral sil-
houette, whole complex forms a rectangle elon-

gated antero-posteriorly and extended somewhat
at each corner. Like the dorsal tubercle, the neu-
ral complex is set slightly to the right of the dorsal
midline.

BRANCHIAL SAC (PHARYNX) (Fig. 5).— Folds
absent, perhaps represented by internal longi-
tudinal branchial vessels. In dissection, 4 inter-
nal longitudinal vessels are evident on each side
of the pharynx; in transverse serial sections, a
fifth internal longitudinal vessel is sometimes
discernible on each side close to the endostyle,
and in a few specimens even a sixth vessel on
each side may run only some length of the sac.
Usually 9 or 10 stigmata lie between these in-
ternal longitudinal vessels. Ten to 1 2 transverse
vessels separate the rows of longitudinally ori-
ented stigmata, and there are about 1 0 parastig-
matic vessels partly or entirely traversing each
side of the pharynx. Along the ventral midline
the branchial sac connects with the body wall by
widely spaced sub-endostylar vascular trabecu-
lae, not by a continuous sub-endostylar mem-
brane. Other vascular trabeculae connect the
branchial sac abundantly in all directions to the

NEWBERRY: DENDRODOA (STYELOPSIS) ABBOTTI, SP. NOV.

243

FIGURE 4. Dorsal tubercle in relation to peripharyngeal groove and dorsal lamina.

FIGURE 5. Right side of pharynx, showing several rows of stigmata and the four internal longitudinal branchial vessels of
the pharynx's right side. Dorsal lamina at top, endostylar groove at bottom. Drawing based in part on unpublished notes of
D. P. Abbott, in part on freshly dissected specimens.

atrial surface of the mantle and to the atrial ep-
ithelium around the gut.

DORSAL LAMINA. — Prominent, continuous,
smooth-bordered dorsal lamina, without lan-
guets.

GUT (Figs. 2, 3). — Esophageal aperture far
dorso-posterior in pharynx; stout esophagus
bends sharply ventrally into stomach; stomach
empties anteriorly into fore-intestine, which
bends to left and passes posteriad on the lateral
side of the stomach. Hind-intestine then curves
sharply dorsad and follows the left mantle to the
anus, which lies slightly to the left-posterior of
the atrial siphon. Stomach has 16 to 18 mod-
erately evident external folds corresponding to
well-developed internal gastric septa. The gastric
septa are reduced to low ridges in the left pyloric
region of the stomach, near the pyloric caecum.
Pyloric caecum is small, sometimes absent. A
highly vascularized pyloric duct joins the sinus-
oidal sheath surrounding the stomach with that
surrounding the fore-intestine. Intestine com-
prises a fore-intestine with a large typhlosole-like
longitudinal plication of its wall and a thick si-
nusoidal jacket between the gut wall and its sheath

of atrial epithelium, and a hind-intestine of more
simply tubular section whose atrial sheath is much
closer to the gut wall. Anus lies dorso-medial or
slightly to the left, above the stomach; anus is
cut square to the axis of the rectum; anal margin
is scalloped into usually 5 lobes that fit together
when the anus is tightly closed.

HEART.— Fairly straight within a somewhat
curved and inflated pericardium; set at about 45°
obliquely to the endostyle, oriented right-ante-
rior to left-posterior, centered roughly beneath
the endostyle in the posterior half of the zooid
(site and orientation in Fig. 2).

OVARY (Figs. 2, 3). — Single, unbranched, sau-
sage-shaped ovary along the right ventral margin
of the zooid, extending almost the entire length
of the zooid, curving sharply dorsad posteriorly
and following the right mantle to arch halfway
over the atrium, recurving dorsally to terminate
in an oviduct directed posteriorly toward the
brood pouch and away from the atrial siphon.
Oviduct lies lateral (away from atrium) to main
mass of ovary, with its lumen penetrating among
the ripening gametes; lateral surface (away from
germinal tissue) of oviduct heavily ciliated, other

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 15

oviducal surfaces apparently not ciliated. Ova-
ries of all specimens examined by dissection or
serial section show all stages of o vogenesis pres-
ent, regardless of season.

TESTIS (Figs. 2, 3).— A dozen to more than 20
separate lobate sacs, not encapsulated with the
ovary but instead lying in the mantle wall ad-
jacent to but clearly outside the ovary's delim-
iting membrane. Most sacs lie medial to the ovary;
some lie anterior to the ovary; few are lateral;
many lobes lie partly "beneath" the ovary, in the
mantle between the ovary and the ventral surface
of the zooid. In many specimens, but not all, a
few testis-lobes lie far posteriorly and on the left
side of the ventral midline, but their spermiducts
join the vas deferens of the testis-lobes that lie
beside the ovary. All sacs join by vasa efferentia
to a single, long vas deferens that lies between
the ovary and the atrial epithelium. This duct
follows the ovary to the region of the atrial si-
phon, and there leaves the ovarian surface to
project toward the atrial siphon from the dorsal
roof of the atrium while the ovary bends pos-
teriad toward its ovipore. The ciliated vasa ef-
ferentia are extremely thin, visible only in serial
section; the vas deferens, also scarcely visible
except in serial section, is a compressed, ciliated
channel terminating in a tiny, spermipore-bear-
ing papilla pointing toward the atrial aperture.
In all specimens examined from all seasons for
gametic condition, many testis-lobes have tailed
sperm, but the spermiducts contain only scat-
tered sperm.

BROOD CHAMBER AND BROODED YOUNG (Figs.
1, 3).— The posterior region of the atrium serves
as a brood chamber, occluded anteriorly by the
branchial sac, on the left by the gut-loop, on the
right by the ascending limb of the ovary. All
specimens examined were brooding young in all
stages of development from (relatively rarely)
fertilized eggs and cleavage stages to (usually)
tadpoles that were still curled (although many of
these straightened upon removal from the brood
chamber during dissections). Quantities of
brooded young vary greatly— fewest (20 to 30)
in midwinter specimens, most (100 to 200) in
late spring to midfall specimens. The brood
chamber often is so swollen with young that it
is readily apparent in living animals. Young are
crammed tightly into the chamber; external study
of zooids divested of test may suggest only a few
larvae, but dissection then reveals many dozens.

The brood chamber is criss-crossed by many vas-
cular trabeculae that connect the branchial sac
and the atrial and gut wall and may keep loose
young from being swept from the brood chamber
by atrial water-currents.

ECOLOGICAL DISTRIBUTION IN CALIFORNIA.—
Intertidal. At Point Pinos and Pigeon Point, peak
abundance is at about +0.3 m, and I have found
no specimens at either site above +0.6 m or
below 0.0 m. This is an open-coast ascidian in
California, inhabiting surf-swept rocky habitats
where the full force of the waves is broken by
surrounding rocks and reefs. Zooids congregate
on horizontal undersides of large boulders, usu-
ally well back from the boulders' edges. Such
boulders restrict waterflow underneath, so much
so that at Point Pinos the rocks that harbor Den-
drodoa abbotti may lie partially in sand that by
its odor and color appears to be virtually anoxic.
At Pigeon Point most rocks with this ascidian
are slightly propped up by their neighbors, so
that oxygen remains plentiful in waters perco-
lating or flowing underneath. Large boulders that
do not have D. abbotti on them may shelter
smaller rocks that do. Many rocks that seem ap-
propriate for this species do not harbor speci-
mens. This spotty distribution of aggregated in-
dividuals may indicate a short swimming period
and quick settlement by brooded larvae, or as
yet unclear ecological restrictions on the adults.
At Pigeon Point, other invertebrates found on
surfaces with Dendrodoa abbotti include the
anemone Epiactis prolifera, the polyclad Noto-
plana acticola, the polychaetes Spirorbis and
Salmacina, the barnacle Balanus glandula (and
sometimes Chthamalus dalli), porcelain crabs
such as Petrolisthes, several encrusting bryozo-
ans such as Eurystomella bilabiata, the asteroid
Leptasterias pusilla, and the aplousobranch as-
cidian Aplidium californicum. But none of these
associated invertebrates seems so severely kept
back from the margins of boulders, so cryptic in
its under-rock habitat, as Dendrodoa abbotti.

ECOLOGICAL DISTRIBUTION IN WASHINGTON.—
Subtidal. At Peavine Pass, San Juan Islands,
specimens were dredged from 10 to 12m. The
species has been sought elsewhere in rocky areas,
but only Peavine Pass, which is swept to the
bottom by strong tidal currents, has proved a
reliable site for collecting by this method, and
even there the species is rarely taken. Debris har-
boring Dendrodoa abbotti contains, as well, Bal-

NEWBERRY: DENDRODOA (STYELOPSIS) ABBOTTI, SP. NOV.

245

TABLE 1. DENDRODOA CARNEA AND D. ABBOTTI: CONSISTENT DIFFERENCES.

Feature

Dendrodoa carnea

Dendrodoa abbotti

Color in life
Siphonal apertures
Dorsal tubercle

Transverse branchial
vessels and rows of stig-
mata

Internal longitudinal
branchial vessels and
folds (DL = dorsal
lamina, (#) = number
of vessels in fold, E =
endostyle)

Endocarps
Margin of anus
Ovary

Testis
Spermiduct

Brood chamber
Brooded young

Bright pink to blood red.
Bilabial.

Narrow ovoid slit whose axis is oriented almost
anterior-posterior.

1 7 or more.

Left: DLO(1)0(1)0(1)0(1)OE as in D. abbotti.
Right: DLO(4-5)0(1)0(1)0(1)OE; prominent low
fold carrying at least 4 vessels on right pharyngeal
wall.

Many, small, widely scattered over entire atrial
wall.

"Reflected but not lobed, often somewhat two-
lipped" (van Name 1912, p. 587).

Straight along right ventral margin of body; ovi-
duct continues so.

Not clearly encapsulated with ovary, testis-lobes
extend somewhat into body wall, predominantly
ventro-lateral to ovary; all testis-lobes close to
ovary.

(?) as in D. grossularia, many short spermiducts
converge in multiple spermipores on atrial sur-
face of ovary (?)

Extensive, including right-posterior region be-
yond oviduct there.

(From a small sample) only a few dozen embryos
brooded at a time.

Gray to ochre, occasionally reddish around si-
phonal apertures.

Bilabial, somewhat more pronouncedly so than
in D. carnea.

Fairly sharply bent "C" whose long axis is ori-
ented laterally.

Ca. 12.

Left: DLO(1)0(1)0(1)0(1)OE as in D. carnea.
Right: DLO( 1)0(1)0(1 XX 1)OE; no multi-vessel fold
on right pharyngeal wall.

Fewer, larger, more (but not entirely) confined to
ventral atrial surface.

Scalloped into usually 5 lobes.

Along right-anterior ventral margin of body, then
bends sharply into ascending limb, recurves be-
hind atrial siphon into dorsal oviduct that pro-
jects posteriorly.

Clearly not encapsulated with ovary, testis-lobes
lie wholly in body wall, predominantly ventro-
medial to ovary; often one posterior group of
testis-lobes far from ovary.

Single, long vas deferens on atrial surface of ovary
receives vasa efterentia of all testis-lobes, ends
mid-dorsally in spermipore-bearing papilla
pointing at atrial siphon.

More restricted to far posterior part of body.

Many dozens to more than 1 00 embryos brooded
at a time.

anus nubilis (one of the best indicators that the
ascidian may be present) and the hydrocoral Al-
lopora. The ascidian occurs especially around the
husks of dead barnacles and in crannies in large
rocks. But dredging of course destroys the set of
surfaces and actual relationships among mem-
bers of the fauna at the site, and so no compar-
ison can yet be made between the subtidal hab-
itat of Dendrodoa abbotti at Peavine Pass and its
intertidal circumstances at Pigeon Point.

The bathymetric contrast between California
and Washington (San Juan Islands) records of
Dendrodoa abbotti is striking. The species may
occur subtidally in California; its inaccessibility,
beneath large boulders, could account for the cur-

rent lack of such records by dredging or even by
diving. But D. abbotti does not occur in the very
low intertidal zone in California, below about
mean low-low tidal levels. Thus, if it does occur
subtidally, there is not a continuous distribution
of the species from those depths to the low- to
mid-tidal habitats where it characteristically is
found. In contrast, in the San Juan Islands, I have
not found the species at all intertidally in habitats
that resemble California's coastal sites— except,
of course, for the lack of surf in the San Juans.
Dendrodoa abbotti appears to be only a subtidal
species in that archipelago. But to the west of the
San Juan Islands, on the southwest coast of Van-
couver Island, B.C., Dr. Ivan Goodbody has

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 15

found this species "on the underside of boulders
at extreme low tide ... on the open coast north
of Sooke." Dr. Goodbody reports (pers. comm.)
that the site there is "a rough boulder strewn
shore with many large rounded boulders indi-
cating heavy wave action." His record thus ex-
tends the intertidal range of D, abbotti into those
Canadian habitats where surf does resemble Cal-
ifornia's. Dr. Goodbody's Canadian specimens
of D. abbotti are now in the collection of the
California Academy of Sciences.

DISCUSSION

A. Comparison of Dendrodoa abbotti
with D. carnea

A comparison of Dendrodoa abbotti with the
western North Atlantic species D. carnea— the
styelopsid dendrodoan that most closely resem-
bles D. abbotti— indicates an array of differences,
some trivial, some marked, but all consistent.
This comparison sets a great many specimens of
D. abbotti against necessarily only a few dissected
specimens of D. carnea (from the USNM col-
lection) and others' reports on D. carnea (see van
Name 1912, 1945). But differences that emerge
even in this perhaps unbalanced sampling of these
species gain force as they become elements in a
consistent pattern of distinctions between the two
taxa, and this pattern has become more persua-
sive with each examination of new specimens.
Table 1 summarizes the comparison.

Even arguably minor distinctions (for exam-
ple, the character of the brood chamber or of the
endocarps) take on significance in Table 1's ar-
ray. The two species are most effectively distin-
guished, however, by the following criteria:

1 . shape and disposition of the ovary;

2. testis-ovary relationship, including D. abbot-
ti's posterior group of testis-lobes;

3. structure of the spermiducts, especially of the
vas deferens;

4. arrangement and number of internal longi-
tudinal vessels of the right side of the pharynx;

5. number of transverse branchial vessels and
rows of stigmata on both pharyngeal walls;

6. shape and orientation of the dorsal tubercle;

7. color in life.

B. Generic Traits

By most accounts and diagnoses, in the genus
Dendrodoa the testis and ovary are "encapsu-

lated" within a common sheath (Monniot and
Monniot 1972), and the testis comprises many
lobes that do not lie in the body wall but rather
hug the parietal (away from the atrium) surface
of the ovary (Huntsman 1913). Most of the go-
nad of Dendrodoa grossularia shows this con-
dition of encapsulation and testis-ovary juxta-
position clearly, although some of the anterior
testis-lobes do lie more in the body wall than
wholly against the ovary. Dendrodoa carnea ex-
hibits a somewhat looser gonadal arrangement:
the testis-lobes apparently are still encapsulated
with the ovary and lie largely against its parietal
surface, but they extend into the adjacent body
wall much more than do the testis-lobes of D.
grossularia, especially to the lateral (right) side
of the ovary. Dendrodoa abbotti carries this loos-
ening of the testis-ovary bond further still: the
testis-lobes of D. abbotti lie "beneath" the ovary
or close by on the medial (left) side of the ovary,
but they lie in the body wall itself, not against
the ovarian mass, and there is no sheath enclos-
ing these gonadal elements into a single structural
unit of intimately juxtaposed parts. And al-
though most of the testis-lobes of D. abbotti lie
very close to the ovary, there is often a group of
testis-lobes lying in the far posterior atrial floor
of the zooid, and actually on the left side of the
zooid, although even this separated and isolated
group is still connected by a vas deferens to the
common spermiduct of all the other, "ovary-
affiliated" testis-lobes.

Dendrodoa carnea is so much like D. grossu-
laria (Traustedt's (1883) type species of his genus
Styelopsis, now a subgenus of Dendrodoa} that
Arnback (1922) and Hartmeyer ( 1 903) have both
suggested these could be merely geographic vari-
ants of a single species— a view not held, how-
ever, by van Name (1945). The main distinction
between these two species is their different num-
ber of internal longitudinal branchial vessels,
more numerous in D. grossularia than in D. car-
nea. But the slight gonadal contrast reported here
also seems to be a consistent one. The difference
takes on added taxonomic significance when D.
abbotti joins the comparison, because the genus
thereby shows a series of testis-ovary juxtapo-
sitions from a tightly joined one to an appreci-
ably looser one— from the condition "character-
istic" of the genus Dendrodoa to one rather akin
to that of the genus Styela.

Perhaps the perplexing Dendrodoa uniplicata

NEWBERRY: DENDRODOA (STYELOPSIS) ABBOTTI, SP. NOV.

247

Hartmeyer 1903, which Millar (1966) redesig-
nates Styela uniplicata Bonnevie 1896 because
"the structure of the gonad agrees better with
Styela," extends the grossularia-carnea-abbotti
series of gonadal arrangements further while re-
taining dendrodoan features of the pharnyx. Un-
fortunately, the meager remnants currently
available of Dendrodoa (or Styela) uniplicata will
not by themselves resolve this question.

Another dendrodoan trait from which Den-
drodoa abbotti diverges involves the spermiduct.
In the genus Dendrodoa, testis-lobes empty in
groups into very short vasa deferentia or even
more cloaca-like pits on the atrial surface of the
ovary, and there are several such spermiporal
loci on the ovary (Berrill 1950). The repetition
of short vasa deferentia, each emptying a group
of testis-lobes, is not usually as striking in D.
grossularia as in the somewhat stylized depiction
of this trait by Lacaze-Duthiers and Delage
(1892), from which work many accounts of the
species have been partly drawn. But Riedlinger
(1902) indicates in his careful study how slight
or even absent the vasa deferentia may be in that
species, in place of which spermiporal loci serve
the converging vasa efferentia of groups of testis-
lobes. Dendrodoa carnea also appears to have
multiple spermipores along the atrial surface of
the ovary (again, though, a condition difficult to
discern in dissections). In contrast, the gonad of
D. abbotti has a single, long vas deferens, as in
Cnemidocarpa and Styela (Fig. 2). All the sper-
miducts of this species are exceedingly fine, and
their disposition difficult to trace except in serial
sections. Such a close scrutiny of D. carnea would
seem appropriate, to find out if that species is
intermediate between D. grossularia and D. ab-
botti in this trait, as it is in testis-ovary juxta-
positions.

Dendrodoa (Styelopsis) abbotti is placed in
Dendrodoa by its possession of a single gonad,
and in Styelopsis because of its unbranched ovary
and its simple pharynx, which lacks folds and
possesses few internal longitudinal vessels. Den-
drodoa abbotti is so much like D. carnea, which
in turn is so much like D. grossularia, that this
placement of the new species seems indisputable.
But the consequence is to relax and modify long-
held gonadal criteria of Dendrodoa, recognizing
that species with styelan gonadal patterns or
cnemidocarpan spermiducal patterns occur in the
genus.

ACKNOWLEDGMENTS

A grant from the Faculty Research Committee
of the Academic Senate of the University of Cal-
ifornia, Santa Cruz has supported much of the
research reported in this paper. I appreciate the
assistance of Linda Cole, U.S. National Museum
of Natural History, who guided me through the
collection there, with the consequence that Den-
drodoa carnea came into consideration at a crit-
ical moment in this study. Professor Ivan Good-
body has shared with both Professor Abbott and
me several Canadian specimens of Dendrodoa
abbotti and ecological information about their
site; I am grateful for his help and for his read-
iness to include this important northern inter-
tidal record in this initial paper about the new
species. Donald P. Abbott, without realizing at
the time the nomenclatural consequence of his
generosity, shared with me his notes and draw-
ings of many years' acquaintance with the species
described in this paper, and I am most grateful
for these and for many other ways in which he
has encouraged me.

LITERATURE CITED

ARNBACK-CHRISTIE-LINDE, A. 1922. Northern and arctic in-
vertebrates in the collection of the Swedish State Museum.
8. Tunicata. 1 . Styelidae and Polyzoidae. In Kungl. Svenska
Vetenskapsakad. Handlingar 63(2): 1-62, pis. 1-3.

BERRILL, N. J. 1950. The Tunicata, with an account of the
British species. London: Ray Society. 354 pp.

BONNEVIE, K. 1896. Ascidiae simplices og Ascidiae Com-
positae fra Nordhavs Expeditionen. In Norske Nordhavs-
Expedition 23(2): 1-16, pis. 3, 4.

GALIGHER, A. E. AND E. N. KOZLOFF. 1964. Essentials of
practical microtechnique. Philadelphia: Lea & Febiger. 484
pp.

HARTMEYER, R. 1903. Die Ascidien der Arktis. In ROmer,
F. and F. Schaudinn, Fauna Arctica 3(2):91-412, pis. 4-14.

HUNTSMAN, A. G. 1913. The classification of the Styelidae.
Zool.Anz. 41:482-501.

LACAZE-DUTHIERS, H. DE AND Y. DELAGE. 1892. Etudes sur
les ascidies des cotes de France. Faune des Cynthiadees de
Roscoff et des c&tes de Bretagne. M6m. Acad. Sci. France
(ser. 2)45:1-323.

LIGHT, S. F., R. I. SMITH, F. A. PITELKA, D. P. ABBOTT AND
F. M. WEESNER. 1954. Intertidal invertebrates of the cen-
tral California coast. 2nd ed. Berkeley: Univ. Calif. Press.
446 pp.

MILLAR, R. H. 1966. Tunicata Ascidiacea. Marine inverte-
brates of Scandinavia, No. l.Oslo:Universitetsforlaget. 123
pp.

MONNIOT, C. AND F. MoNNioT. 1972. C16 mondiale des gen-
res d'ascidies. Arch. Zool. Exp. Gen. 1 13:31 1-367.

RIEDLINGER, R. 1902. Untersuchungen ilber den Bau von
Styelopsis grossularia des Ostsee. Nova Acta Akad. Leop.-
Carol., Halle 81:1-62, pis. 1-6.

248 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 1 5

SMITH, R. I. AND J. T. CARLTON, eds. 1975. Light's manual: VAN NAME, W. G. 1912. Simple ascidians of the coasts of

intertidal invertebrates of the central California coast. 3rd New England and neighboring British provinces. Proc. Bos-

ed. Berkeley: Univ. Calif. Press. 716 pp. ton Soc. Nat. Hist. 34:439-619, pis. 43-73.

TRAUSTEDT, M. P. A. 1883. Vestindiske ascidiae simplices. . 1945. The North and South American ascidians.

2. Molgulidae og Cynthiadae. Vid. Medd. Naturhist. Kj5- Bull. Amer. Mus. Nat. Hist. 84:1-476, pis. 1-31.
benh., ann. 1882:108-136, pis. 5, 6.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 16, pp. 249-267, 6 figs., 6 tables

December 11, 1984

THREE NEW SPECIES OF SEVEN-GILLED HAGFISHES

(MYXINIDAE, EPTATRETUS) FROM

THE PACIFIC OCEAN

By
Charmion B. McMillan and Robert L. Wisner

Marine Biology Research Division, A -01) 2, Scripps Institution of Oceanography,
La Jolla, California 92093

ABSTRACT: Three new species of hagfishes (Myxinidae, Eptatretus) from the Pacific Ocean are described,
and compared with /-.. cirrhatus. All four species have seven pairs of gill pouches and associated external
openings. Of the new species, /:. carlhubbsi is known from Molokai to Guam, north-central Pacific, K.
laurahubbsi from off south-central Chile, and /:. strahani from near Lubang Island, Philippines, South China
Sea. Eptatretus cirrhatus occurs in the Australian-New Zealand area. Methods used in examination of hag-
fishes are described, and sensory (lateral line) canals are delineated and discussed briefly.

INTRODUCTION

This study of seven-gilled hagfishes (genus Ep-
tatretus) from the Pacific Ocean is one of a series
resulting largely from the specimens and data
accumulated under direction of the late Carl L.
Hubbs. Herein we describe three new species,
present new data on E. cirrhatus (Bloch and
Schneider 1801), offer suggestions for initial
preservation of myxinids to provide good study
material, and discuss methods useful in the taxo-
nomic study of hagfishes. We also offer figures
and a brief description of the sensory canals found
in the ocular regions of two of the four species.

DISCUSSION

Our examinations have shown that species of
Eptatretus from the Pacific Ocean have six to
fifteen pairs of gill pouches and corresponding
external apertures. The three new species de-
scribed below, with Eptatretus cirrhatus, com-
prise a group having seven pairs of gill pouches.

One aberrant specimen has eight pouches on each
side, but with corresponding apertures arranged
abnormally. Our rather limited counts (22 pairs)
from the three new species may not reflect ex-
tremes of variation, but the number of gill ap-
ertures in Eptatretus cirrhatus appears to be con-
stant—seven pairs in 48 specimens. In 44 counts
from 22 specimens of the three new species, the
only variation from seven was the specimen cited
above (further discussed and figured below).

Counts of six apertures for Eptatretus cirrhatus
recorded in the literature apparently resulted from
a confusion of species. Giinther (1870) stated
that the species had "six or seven gill openings
on each side," but he listed specimens from South
Africa (E. hexatrema Miiller, 1834) and Japan
(E. burgeri Temminck and Schlegel, 1850).
Species from these areas commonly have six pairs
of gill openings. Referring to Eptatretus cirrha-
tus, Waite (1909) stated, "The gill-openings ap-
pear to be seven in number, but I have seen an

[249]

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16

example in which there were but six openings on
one side, though seven were present on the other."
A variation of one per side is common in species
having ten or more gill pouches, such as E. deani
(Evermann and Goldsborough 1907) and E.
stoutii (Lockington 1 878). Also, it is possible that
Waite had an abnormal specimen in which two
pouches shared the same opening (see above).
Strahan's (1975) finding of "seven (rarely six)
pairs of branchial apertures" for Eptatretus cir-
rhatus may have been influenced by Giinther's
or Waite's accounts.

Regan (1912) listed a species with "7 gill open-
ings: on each side two rows of 8 teeth. Southern
Pacific" as Heptatretus banksii, and placed in its
synonymy Homea banksii Fleming 1822, and
Bdellostoma heptatrema Miiller (1834). Regan's
total count of 32 teeth is much lower than that
of any of the four species treated herein (Table
6), and may indicate an erroneous count or an
undescribed species. Regan may have counted
three fused median teeth (multicusps) on each
row as one, thus reducing the count to 32 from
a possible 40. This would have been much nearer
our minimal count of 43 for Eptatretus cirrhatus,
under which we synonymize the above three
names.

Species of Eptatretus having seven gill aper-
tures are not restricted to the Pacific Ocean. Fern-
holm and Hubbs (1981) listed a species having
seven apertures from the Caribbean Sea. Fern-
holm (1982) has further described it as new.

In general, we concur with Fernholm and
Hubbs in terminology, with but minor varia-
tions. We believe the term "dental muscle" is
more appropriate than "tongue," "lingual," or
"club-shaped muscle" in reference to the firm
elongate complex of muscles and cartilages which
constitutes the feeding mechanism of myxinids.
Apparently the term "tongue" was first used by
Miiller (1834), but we concur with Ayers and
Jackson (1900) that the entire apparatus in no
way resembles a tongue. They stated, "The ho-
mology of this organ with the vertebrate tongue
has never been discussed, nor do we know of any
effort to determine the true nature of this organ."
Dawson (1963:248, fig. 11) provided a detailed
analysis and figure of the structure, and of the
"teeth" and "jaw apparatus." She concluded (p.
253) that it was unwise to make any definite
assumptions concerning homologies of the car-
tilages and muscles.

There are two pairs of anterior and posterior
sets (series) of sharply pointed, laterally flattened,
horny structures in the oral cavity which are
embedded in a dental plate. These structures cut
and scrape food into ingestible portions when
everted and retracted by the dental muscle. Al-
though the term "teeth" has been widely used in
reference to these structures, they are unlike the
teeth of other vertebrates, being composed en-
tirely of keratin and devoid of calcification. Daw-
son (1963:247) concluded that, "It is most likely
that there is no phylogenetic connection between
these teeth and calcified teeth, and that they are
an individual adaptation to a parasitic mode of
life."1 For descriptive and statistical purposes,
we prefer the terms unicusps and multicusps to
differentiate between single and composite teeth—
the latter formed by the fusion of two or three
unicusps. We consider the number and arrange-
ments of both the multicusps and unicusps to be
a significant species character.

MATERIALS

Collection data and disposition of specimens
examined in this study are listed in the treatment
of each species. Institutions which have fur-
nished study material, or in which type speci-
mens have been deposited, are: Bernice P. Bish-
op Museum, Honolulu, Hawaii (BPBM); United
States National Museum, Washington, D.C.
(USNM); Scripps Institution of Oceanography,
La Jolla, California (SIO); California Academy
of Sciences, San Francisco (CAS); Museum Na-
tional d'Histoire Naturelle, Paris, (MNHN);
University of the Philippines Zoological Mu-
seum, Diliman, Quezon City, Philippines
(UPZM); Australian Museum, Sydney (AMS);
Zoological Institute, Academy of Sciences, Len-
ingrad (ZIN).

METHODS

The methods of measuring and counting de-
scribed herein represent original methods as well
as some used by prior authors including Dean
(1904), Nani and Gneri (1951), Richardson
(1953), and Strahan (1975). Fernholm and Hubbs
(1981) reported many of these methods in their
study of the eastern Atlantic Eptatretus. When

1 Hagfishes are not parasitic; they scavenge dead or mori-
bund fishes and invertebrates.

MCMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES

251

the senior author, in collaboration with the late
Carl L. Hubbs, began work on the myxinids (in
1969), it was obvious that no standard criteria
existed for the study of hagfishes, which lack the
jaws, opercula, rayed fins, scales, gill rakers, and
bones found in most fishes. Early workers ap-
plied different names to the same anatomical
characters, defining them differently or not at all,
and often not mentioning the methods used in
measuring and counting. Therefore, it was dif-
ficult to correlate or compare data of different
authors, and taxonomic confusion resulted. We
hope that the methods proposed and defined be-
low will provide future investigators with a stan-
dard by which hagfish species and specimens may
be readily compared and identified.

Proper treatment immediately after capture is
of particular importance in rendering specimens
suitable for study. Often too many live hagfish
are crowded in jars of preservative, resulting in
coiled or bent bodies, usually heavily coated with
slime (mucus) and difficult to measure or count.
The copious secretion of slime, characteristic of
the family Myxinidae, is dramatically curtailed
by prompt immersion in fresh water, preferably
warm. This rapidly kills the hagfish and prevents
further extrusion of slime, which otherwise con-
tinues for several minutes even in formalin. Any
remaining slime may be removed with paper or
cloth towels, and the specimens should then be
laid straight in a suitably large container of for-
malin until fixed. If a specimen is too large for
a flat pan, it should be coiled smoothly in a 3-
5 -gallon container, taking care not to deform the
snout or twist the body, and covered with for-
malin. This treatment produces fairly straight
specimens with a minimal coating of slime, and
greatly facilitates accurate counts and measure-
ments.

Since fresh hagfishes deteriorate rapidly, pres-
ervation should be prompt. Color photos or notes
should be made to record pigmentation, and tis-
sue or blood desired for biochemical or chro-
mosomal studies should be taken prior to im-
mersion in formalin. We find that initial freezing
prior to chemical preservation may cause soft-
ening of the tissue and collapse of eggs and in-
ternal organs, but it may be preferable to crowd-
ing into a too-small container. Due to the many
body openings, we consider it unnecessary to slit
the skin or to inject preservatives; hagfishes are
so soft that the skin may tear and some under-

lying tissues may come apart, causing difficulty
in subsequent measures and counts.

ABBREVIATIONS

PCD: external opening of the pharyngocuta-
neous duct; ordinarily confluent with the pos-
teriormost left gill aperture, and much larger than
all other apertures.

GA: gill (branchial) aperture; external opening
of the efferent duct leading from a gill pouch.

GP: gill pouch; rounded, serially arranged
structures along and posterior to the dental mus-
cle.

DM: dental muscle; the firm, elongate, cylin-
drical complex of muscles and cartilages that
moves the dental plates and sets of cusps during
feeding. Posterior portions of DM are shown in
Figure 3.

VA: ventral aorta; the portion between the heart
(ventricle) and where it branches to each side of
DM.

ABA: afferent branchial artery; one of the small
blood vessels that lead to each gill pouch from
VA or its branches.

MEASUREMENTS

If the specimen is distorted due to preserva-
tion, it should be moderately straightened to ap-
proximate its normal form. Measurements are
taken from the left side with the fish lying on a
meter stick; dividers or dial calipers are advisable
for shorter lengths. We arbitrarily divided the
body into four major sections (Fig. 1):
prebranchical, branchial, trunk, and caudal. These
are particularly apropos to genera Eptatretus and
Paramyxine, as each has more than one GA, thus
a branchial section. In Myxine, Neomyxine, and
Nemamyxine, there is only one GA on each side,
that on the left being confluent with PCD.

Synonymous terms appearing in the literature
are: "head" or "pectoral" for prebranchial, "gill"
for branchial, and "abdominal" for trunk. The
term "mucus" has often been used for slime,
"teeth" for cusps, "tongue" or "lingual muscle"
for dental muscle, and "outer" and "inner" for
posterior and anterior in referring to the series
of cusps.

Body measurements we have found particu-
larly useful are:

Total length (TL): snout (anterior tip of ros-
trum, excluding barbels) to posteriormost mar-
gin of tail or caudal fin.

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16

FIGURE 1. A-B: Sketches of an Eptatretus and a Myxine, showing regions of body used in study of myxinids: 1 to 5, total
length; 1 to 2, prebranchial; 2 to 3, branchial; 3 to 4, trunk; 4 to 5, caudal. C: sketch of head region of a myxinid, showing barbel
pairs 1 , 2, and 3, and nasopharyngeal opening, 4.

Preocular length: snout to center of eyespot,
unpigmented area (if present) marking the ocular
region.

Prebranchial length: snout to front of first, or
only, GA.

Branchial length: front of first to front of last
GA (PCD). The anterior edge of the last GA is
used because the posterior margin is often too
vague and poorly defined to provide a definite
reference point.

Trunk length: front of PCD to origin of cloaca.

Body width: maximum dimension about mid-
way between rostrum and PCD.

Body depth: maximum vertical depth in trunk
region, including finfold if present; depth ex-
cluding finfold should be taken at the same place.
In both width and depth measurements the body
should be molded into a seemingly natural shape
if necessary.

Depth at cloaca: vertical depth at origin of
cloaca.

Tail depth: maximum vertical depth of flat-
tened tail, with any roll-up or fold of the thin
tail margin uncurled and flattened.

Barbel length: from center of base to tip of
each barbel (Fig. 1). The distance between bases
of each pair may be measured from the inside
edge of each base. Barbels are often curled and
difficult to measure accurately, but in certain
species barbel length may be a significant char-
acter, and is worth measuring.

Dental muscle length (DM): snout to tip of
DM, as revealed by a midventral incision in the
prebranchial region.

Dental muscle width: measured at a straight-
sided portion well anterior to tapering end.

Dental muscle depth: measured at same place
as width measure. Rather than using the total

MCMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES

253

length, we have found it convenient to compare
the length (or width) with the unbranched por-
tion of the VA with measurements of the DM.
This is a significant ratio in certain species, but
varies greatly between specimens of other species.
Weight: may be taken, but we have not found
it to be a reliable or useful character, principally
because of the uncertainty in determining if all
the entrapped fluid was drained, and because of
dehydration of body fluids during preservation.

COUNTS

Ordinarily the branchial openings (GA) are the
first items examined to ascertain the genus and
possible species. The gill pouches are usually
counted after the teeth (cusps) when the oral cav-
ity incision is extended midventrally to the re-
gion of the PCD. Before counting the slime pores,
we gently scraped away any coagulated slime
overlying the line of pores; an air jet greatly fa-
cilitated location of pores. Because so few spec-
imens were available for this study, both sides
were counted to obtain wider range of variation.
Counts we have found particularly useful are:

Slime pores:

Prebranchial— from anteriormost slime pore
to last one before first GA.

Branchial— those pores in immediate associ-
ation with (usually below and to the right of)
each GA; often one less than GA count in Ep-
tatretus, and much less, or absent entirely, in
Paramyxine. There is usually no slime pore as-
sociated with PCD, but this varies with species
and individual specimens. In this study all species
except E. strahani have a branchial pore count
equal to or higher than the number of GA; the
extra pores vary in location and number.

Trunk— the series posterior to PCD and ter-
minating anterior to end of cloaca, distinctly sep-
arate from cloacal series.

Cloacal— the pores distinctly before a vertical
from posterior end of cloaca, usually starting
somewhat anterior to and elevated from origin
of cloaca.

Caudal— from first pore distinctly behind a
vertical from posterior end of cloaca to last pore
on tail. For statistical purposes we combine counts
of cloacal and caudal pores under the heading
"tail pores" (Table 2).

Cusps (teeth): We refer to a single "tooth" as
a cusp, or unicusp, if it is not fused to one or
more adjoining cusps. A unit of two or more
cusps fused together at some point prior to its

FIGURE 2. Cusps and basal plates, in excised and spread
condition, ofE. carlhubbsi, paratype USNM 233742, 955 mm
TL.

embedment in the cartilaginous dental plate is a
multicusp.

The two paired sets of cusps (the outer and
inner rows of Fernholm and Hubbs [1981] and
Fernholm [1982]) are examined from the ventral
aspect. They are revealed by a midline incision
from the base of the oral cavity through the car-
tilaginous pharynx until the sets are free and eas-
ily turned outward for viewing. There are dis-
advantages to this method. It is easy to misjudge
the midline (if the "face" has been distorted in
preservation) and cut through the median teeth,
making counts difficult; also, the resulting view
presented to the observer is a reversed image of
the actual arrangement. The inner left row ap-
pears on the outer right side and vice versa. To
avoid this confusion, the incision may be made
from either side of the oral cavity to just under
the third barbel, then extended laterally down-
ward through the thin membrane, exposing the
paired sets of cusps which, when spread apart,
appear as shown in Figure 2.

On most specimens the count of multicusps
may be determined by placing a dissecting or air
jet needle under the first two cusps and gently

254

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16

I br. VA

FIGURE 3. Ventral view of branchial region of: 1, E. carlhubbsi: 2-3, E. laurahubbsi, showing diversity in afferent branchial
arteries (ABA) leading off from branches of ventral aorta (VA); 4, E. strahani; 5, E. cirrhatus.

lifting; the multicusp usually lifts and separates
from the adjacent unfused cusp. However, in the
nine largest specimens examined by us (E. carl-
hubbsi), lifting often raised the entire dental plate
and set of cusps. Even if cusps are unquestion-
ably fused, a line may extend among the fusion
to the plate or "gum line;" in such instances per-
haps the only valid criterion for separating mul-
ticusps from unicusps is the distinctness of this
line as seen under magnification. Such lines are

in marked contrast to the condition shown by
scanning electron microscopy of E. springeri
(Fernholm and Hubbs 1981: fig. 2), wherein no
lines are evident in the multicusps.

Gill apertures and pouches: In genera Myxine,
Neomyxine, Nemamyxine, and Notomyxine,
dissection is necessary to determine the number
of gill pouches, since only one pair of efferent
ducts leads to the exterior. A midventral incision
is made from the single pair of GA anteriorly

MCMILLAN AND WISNER.: NEW SPECIES OF PACIFIC HAGFISHES

255

until all pouches are revealed (Fig. 3). The cut
should be deep enough to expose VA and ven-
tricle, taking care not to sever branches of VA
or any ABA, or to destroy the origin of the ventral
finfold if it is present anterior to PCD.

There are multiple, readily visible GA in gen-
era Eptatretus (5-15 pairs) and Paramyxine (5-
7 pairs). Although the number of internal pouch-
es ordinarily is the same as the external apertures,
there may be variation; thus, it is desirable to
count the pouches and examine the arrangement
of the GP relative to DM and branched and un-
branched portions of VA (Fig. 3). The arrange-
ment is often of taxonomic importance, although
variation occurs (see E. laurahubbsi).

Sensory canals (lateral lines): Ayers and Wor-
thington (1907:331, figs. 5-10), in a study of the
skin-end organs of the trigeminal and lateralis
nerves of Bdellostoma dombeyi (=Eptatretus
stoutii [Lockington 1 878]), described and figured
lateral line canals, associated dermal grooves, and
nerve endings. They showed the canals as short
lines occurring only dorsally and somewhat lat-
erally on the "head" and in two groups, one be-
fore and one behind the eyespots. Plate (1924:
66, fig. 6 1 D) accepted the interpretation by Ayers
and Worthington that the short lines constituted
lateral line canals, but considered the dermal
grooves to be artifacts. Ross (1963:155) cited
both these studies and stated that the lateral lines
had not been described in Myxine glutinosa. To
our knowledge these are the only prior references
to lateral line canals of hagfishes.

We concur with Ayers and Worthington that
the canals occur only on the head (in the ocular
area of the prebranchial region). However, they
are lateral only in that a few occur on the side
of the head, with most on the dorsal surface (Fig.
4), and none at all on the rest of the body. As-
suming that the canals are indeed sensory in
function, we prefer the term "sensory" to "lat-
eral." Sensory canals occur in only two of the
four species discussed here (E. strahani and E.
cirrhatus, Fig. 4), but not on all specimens, and
are irregular in number and form. The erratic
occurrence in location and in numbers of canals
is intriguing, as is their total absence in two of
the four species.

Due to the limited number of specimens avail-
able, it is difficult to draw any firm conclusions
regarding the taxonomic value of sensory canals.
Ayers and Worthington (1907) stated that these
canals were difficult to find because they were

E. cirrhatus

E . strahani

FIGURE 4. Sketches (not to scale) of head regions of Ep-
tatretus cirrhatus and E. strahani showing arrangements of
sensory canals. The first two pairs of barbels are omitted.

very small and the surface indications faint, and
that any apparent erratic appearance might be
due to the observer. However, on the specimens
examined by us the canals, when present, were
readily visible under adequate magnification and
lighting, and often by the unaided eye. They ap-
pear as thin lines, about 1-3 mm long, variably
straight or curved (Fig. 4), often very slightly
raised above the skin, and sometimes covered
with a coating of coagulated slime. Histological
examination was not done, nor have we attempt-
ed to observe these canals on unpreserved fishes.
Old, healed scars are often present in areas
occupied by the sensory canals, and elsewhere
on the body, mostly anteriorly. These are iden-
tifiable as shallow depressions, usually wider and
longer than the sensory canals. Many scars occur
singly, but often they are in groups of parallel
lines, the spacing closely resembling that of the
anterior cusps. Possibly this scarring occurs when
many hagfishes are feeding in close proximity
competing for food, or when crowded in a trap.

256

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16

TABLE 1 . AVERAGES AND RANGES (IN THOUSANDTHS OF TOTAL LENGTH) OF SELECTED BODY PROPORTIONS FOR FOUR SPECIES
OF SEVEN-GILLED HAGFISHES (GENUS EPTATRETUS) FROM THE PACIFIC OCEAN.

E. carlhubbsi

E. laurahubbsi

E. strahani

E. cirrhatus

N (size range in mm)

9(813-1160)
Avg. (range)

8 (240-375)
Avg. (range)

5 (265-520)
Avg. (range)

8(481-655)
Avg. (range)

Preocular length
Prebranchial length
Branchial length
Trunk length
Tail length
Tail depth
Body depth with finfold
Body depth without finfold
Body depth at cloaca

38 (36-54)
184(168-197)
68 (55-77)
602 (577-623)
160(145-176)
97(89-105)
No finfold
93 (78-106)
73 (65-85)

50 (44-59)
193(184-204)
55 (52-59)
561 (545-585)
198(181-213)
90 (82-99)
89 (74-97)
81(73-91)
70(61-80)

63 (57-68)*
220(210-231)
77 (69-83)
521 (500-537)
182(174-202)
117(109-125)
111 (101-117)
98 (94-105)
87 (77-94)

60 (52-67)
225(214-239)
76 (69-89)
550 (525-563)
154(135-168)
83(77-91)
89 (69-102)
88 (69-102)
67 (57-75)

* Due to lack of visible eyespots, the preocular length was taken from center of uncovered pupil.

Waite (1909) placed three adult E. cirrhatus in
a bucket of formalin and observed them savagely
attacking each other. One was bitten at least 1 5
times by the other two.

KEY TO SEVEN-GILLED SPECIES OF Eptatretus
FROM THE PACIFIC OCEAN

la. Slime pores of trunk 60-70, low, well be-
low mid-lateral aspect. Total cusps 61-71.
Eyespots present 2

Ib. Slime pores of trunk 45-53, high, near
mid-lateral aspect. Total cusps 43-53.
Eyespots present or absent 3

2a. Ventral finfold absent. Two (rarely three)
fused cusps on anterior multicusps, three
on the posterior. Eyespots large, promi-
nent E. carlhubbsi n.sp.

2b. Ventral finfold prominent. Two (rarely
three) fused cusps on each of the four mul-
ticusps. Eyespots present

E. laurahubbsi n.sp.

3a. Ventral finfold readily visible. Eyespots
absent. Ventral margin of tail forming a
nearly straight line from cloaca to abrupt
beginning of curve around tail. Anterior
few gill apertures small, slitlike. No pale
rings around slime pores or gill apertures.
Three fused cusps on each of the four mul-
ticusps E. strahani n.sp.

3b. Ventral finfold vestigial. Eyespots present.
Tail margin smoothly ovate. All apertures
rounded. Pale rings around slime pores
and gill apertures. Three fused cusps on
each of the multicusps E. cirrhatus

Eptatretus carlhubbsi new species

HOLOTYPE.— SIO 68-473, mature female, 96 1 mm TL, taken
at 19°18'N, 166°33.5'E, near Wake Island, in a free-vehicle
trap on bottom at 1574 m, 12-13 Sept. 1968.

PARATYPES.— SIO 68-473, female, 810 mm TL, taken with
the holotype; SIO 82-63 (formerly BPBM 27850), female, 1 1 25
mm TL, taken at Brooks Banks, between French Frigate Shoals
and Gardner Pinnacles, Leeward Islands, Hawaii, Nov. 1981,
Mokihana Cruise 81-12, set 35, shrimp trap, depth not given;
BPBM 27848, male, 1 160 mm TL, taken at 12°56'N, 166°22'W,
French Frigate Shoals, Leeward Islands, Hawaii, 7 Nov. 1981,
shrimp trap at 684 m; BPBM 27851, male, 830 mm TL, taken
off the north shore of Molokai Island, Hawaii, 26-27 Dec.
1981, shrimp trap at 659 m; USNM 227440, male, 900 mm
TL, taken at 24°48'N, 167°14'W, R/V Cromwell Cruise 80-05,
Station 57, in a shrimp trap at 835 m; USNM 233742 (formerly
NMFS P-0289), male, 955 mm TL, taken at 14°59'N, 145°13'E,
Esmeralda Bank, Guam, 5-6 April 1981, Cruise Typhoon 81-
01, Station 151, in a shrimp trap at 1061 m; CAS 50705
(formerly BPBM 27847), male, 1064 mm TL, Leeward Islands,
Hawaii, Nov.-Dec. 1981, depth and method of capture not
given; CAS 50706 (formerly BPBM 27849), male, 980 mm
TL, taken at French Frigate Shoals, East Plateau, north side,
Leeward Islands, Hawaii, 19 Nov. 1981, in a shrimp trap at
481 m.

DIAGNOSIS.— A seven-gilled Eptatretus having
no ventral finfold, very large eyespots, two (rarely
three) fused cusps on the anterior multicusps and
three on the posterior.

DESCRIPTION. —Counts: Those of holotype giv-
en first (left and right sides), followed by ranges
for all specimens in parentheses: gill apertures 7,
7 (all); prebranchial slime pores 15, 16 (12-17);
branchial pores 7, 7 (6-8); trunk pores 60, 61
(60-70); cloacal pores 2, 2 (1-3); caudal pores
11, 11 (11-13); tail pores 13, 13 (12-16); total
slime pores 95, 97 (93-1 10). Cusps on anterior
multicusps 2, 2 (rarely 3); posterior multicusps
3, 3 (all); anterior unicusps 16, 16 (15-17); pos-

MCMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES

257

TABLE 2. PREBRANCHIAL, BRANCHIAL, AND TAIL SLIME PORES OF FOUR SPECIES OF SEVEN-GILLED HAGFISHES (GENUS EPTA-
TRETUS) FROM THE PACIFIC OCEAN.

Prebranchial slime pores

E. carlhubbsi 1

5 2

E. laurahubbsi

3 8

E. strahani

2 3

E. cirrhatus

17 17 4

1 43

Branchial slime pores

E. carlhubbsi

E. laurahubbsi

E. strahani

E. cirrhatus

Tail slime pores'

Tail count is the total of the cloacal and caudal slime pores.

E. carlhubbsi

8 1

E. laurahubbsi

12 2

E. strahani

E. cirrhatus

tenor unicusps 12, 13 (11-13); total cusps 68
(64-71).

Morphometry: In thousandths of total length;
values for holotype given first, followed by ranges
for all specimens: preocular length 38 (36-54);
prebranchial length 184 (168-197); branchial
length 68 (55-77); trunk length 602 (577-623);
tail length 160 (145-176); tail depth 97 (89-105);
body depth 90 (78-1 12); depth at cloaca 74 (62-
86).

All specimens from Hawaii were frozen ini-
tially; the body proportions of these may not be
closely comparable to the other collections, which
were initially preserved in formalin. It is not
known what effect freezing may have on subse-
quent shrinkage, but it is possible that the soft
tissues of hagfishes are greatly affected by the

expansion of cells in freezing. It is known that
length of time in preservative significantly affects
the total length; a shrinkage of 10% is not un-
common. However, to our knowledge no study
has been done showing the changes in other body
proportions. Body proportions (Table 1) and
counts (Tables 2-6) are compared with similar
data for other seven-gilled Eptatretus from the
Pacific Ocean.

Body robust; prebranchial region slightly
deeper than wide; body increasingly compressed
laterally to tail, varying in greatest depth from
8% to 1 1% of TL. Two to four GP anterior to
tip of DM, which is somewhat flattened poste-
riorly. Length of DM 19% (17-21%) of TL, its
width 15% (9-19%) of its length, its depth 66%
(57-87%) of its width. VA short, wide, its width

TABLE 3. TRUNK SLIME PORES OF FOUR SPECIES OF SEVEN-GILLED HAGFISHES (GENUS EPTATRETUS) FROM THE PACIFIC OCEAN.

Trunk slime pores

45 46 47 48

49 50 51 52 53 54 55 56 57 58 59 60 61

62 63 64 65 66 67 68 69 70

E. carlhubbsi

1 2

3 145 2

E. laurahubbsi

3 1

22 125

E. strahani

2223

E. cirrhatus

5 2 10

11 3 5 2 5

258

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16

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39% (35-43%) of its length. Distance between
DM and branching of VA 17% (12-22%) of DM
length. GA rather evenly spaced in a line often
sloping or curving slightly downward posteriorly,
with the exception of paratype BPBM 27851,
apparently an aberrant specimen. It has eight GP
on each side, but with corresponding apertures
arranged abnormally (Fig. 6); there are eight GA
on the left side, but only seven on the right, with
two gills sharing one aperture.

Head blunt, face sloping at about 30° from the
vertical. Eyespots large, prominent, ranging from
nearly round to square or rectangular. Removal
of overlying integument shows the embedded eyes
varying from under to well behind center of eye-
spots. Eyes nearly round, about 5 by 6 mm, slant-
ing ventrodorsally at about 45°. The first two
pairs of barbels of nearly equal length on the
holotype, the second pair slightly shorter than
the first. Proportions of the barbel pairs on the
paratypes show great variation, with some of the
first pairs shorter than the second. The second
barbel, right side, of paratype SIO 68-473 is bi-
furcate to the base, both halves of about equal
length. No sensory canals are present on any of
the specimens.

There is no ventral finfold, but an intermittent,
vague line resembling a median suture extends
from well behind PCD nearly to cloaca. The thin
margin of the tail, common on other hagfishes,
is not readily evident in these large specimens.
If a caudal finfold is present, it is very thick,
tapering steeply from the tail musculature. This
may be a species character or merely the result
of age or size; no juveniles are known. In the
other three species discussed below (particularly
the juvenile specimens ofE. laumhubbsi) the tail
margins are notably deeper and thinner. The av-
erage tail length in the nine specimens of E. carl-
hubbsi is about twice (1.9-2.5) the body depth
at the origin of the cloaca, but the shapes of the
tails vary considerably (Fig. 5). The deep notch
on the dorsal surface of the tail of the holotype
is apparently an old, entirely healed injury; no
notch is present on any paratype. The tail depths
average 58% (49-65%) of the lengths; shape var-
ies from that of a truncated club (No. 7) with no
evidence of finfolds to the very deep, expanded
form of No. 3, which has a notably less thickened
margin.

The holotype and three paratypes are tan in
color; two are from Wake Island, and one each

MCMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES

259

TABLE 5. UNICUSPS IN THE ANTERIOR AND POSTERIOR SETS OF CUSPS OF FOUR SPECIES OF SEVEN-GILLED HAGFISHES (GENUS
EPTATRETUS) FROM THE PACIFIC OCEAN.

Anterior unicusps

12 13 14 15

E. carlhubbsi

E. laurahubbsi

1 1 4

E. strahani

E. cirrhatus 1

Posterior unicusps

10 11

14 15 16 N

E. carlhubbsi

E. laurahubbsi

1 16

E. strahani

E. cirrhatus

from French Frigate Shoals and Molokai, Ha-
waii. Five paratypes (one from Guam, four from
Leeward Islands, Hawaii) are a purplish brown;
the Guam specimen has a more brownish over-
tone. Scattered, irregularly shaped pale areas of
varying sizes occur mainly on the anterior por-
tions of the body and occasionally as small patches
on the other parts; these light spots are more
visible on the purplish-brown specimens. Small,
very dark brown blotches occur randomly on all
but one of the tan specimens. Pale rings surround
the GA, but not the slime pores. The color dif-
ferences are striking, but such variation is known
in other species wherein blotching, lighter shades,
and even albinism occur (Dean 1903; Jensen
1959).

The color variations discussed above and the
striking differences in tail shapes and barbel ra-
tios, as well as wide ranges in other body pro-
portions and differences, often indicate the pres-
ence of more than one species, but we find no
definite correlation between these characters and
those of counts, sex, other body proportions, or
geographical areas. Further collections may show

that subspecies or other taxa occur within or be-
tween the three areas— Hawaii, Wake Island, and
Guam.

The first few prebranchial pores curve gently
downward on the holotype, but on all paratypes
they are in an essentially straight line (occasion-
ally the first one or two are slightly depressed).
There are seven or eight slime pores in the bran-
chial region, one or two more than expected,
which is one less than the number of GA.

Eggs: The holotype contained at least 2 1 eggs,
the largest 58.5 by 14.4 mm. These eggs had been
removed and bottled separately by an earlier in-
vestigator, and it is not known that all were pre-
served. None were in the specimen when ex-
amined by us. Paratype BPBM 27848 contained
32 eggs, the largest 75.5 by 16.5 mm. Some eggs
were damaged by freezing, the original preser-
vative, and were loose in the body; these are now
bottled separately. Eight eggs, the largest 58.5 by
14.4 mm, were packaged separately within a larg-
er pack containing a male and a female (SIO 82-
63, formerly BPBM 27850). The female con-
tained only 1 3 eggs, the largest 63.0 by 1 2.6 mm.

TABLE 6. TOTAL CUSPS OF FOUR SPECIES OF SEVEN-GILLED HAGFISHES (GENUS EPTATRETUS) FROM THE PACIFIC OCEAN.

Total cusps

43 44

45 46 47

48 49 50

51 52 53

54 55 56 57 58 59 60 61

62 63

64 65

66 67 68

70 71

E. carlhubbsi

2 1

E. laurahubbsi

E. strahani

E. cirrhatus

1 2

282

3 1 2

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i i i i i I i i i i I
0 10 20 cm

MCMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES

261

•&&:

O O

FIGURE 6. Sketch of abberrant arrangement of gill apertures
of a specimen of E. carlhubbsi, paratype BPBM 27851, 830
mm TL.

Although there was no identifying label with the
separate eggs, we assume they had been taken
from the same female. Since the Hawaiian spec-
imens apparently were all frozen as initial pre-
servative, these eggs could have been stripped
prior to immersion in preservative; the specimen
had not been cut open anywhere on the body.
Although all were extremely large, none of the
eggs cited above had the terminal hooks of fully
ripe eggs (Dean 1899; Jespersen 1975).

ETYMOLOGY.— With great respect and admi-
ration we dedicate this species of giant hagfish
to the late Carl L. Hubbs, himself a giant in ich-
thyology.

Eptatretus laurahubbsi new species

HOLOTYPE.— SIO 65-643, juvenile female, 375 mm TL, tak-
en at 33°31'S, 78°50'W, near Mas a Tierra, Islas Juan Fernan-
dez, in a free vehicle trap on bottom at 2400 meters, between
hours of 2030 and 0830, 12-13 Dec. 1965, Cruise 12 of R/V
Anton Bruun.

PARATYPES (remaining material examined). — Seven juve-
niles (sex questionable) taken with the holotype, are deposited
as follows: SIO 65-643, two, 369 and 287 mm TL (deposited
with the holotype); CAS 49125, two, 287 and 358 mm TL;
USNM 227441, two, 240 and 265 mm TL, Museo Nacional
de Chile, Santiago, one, 240 mm TL.

DIAGNOSIS.— A seven-gilled Eptatretus having
a well-developed finfold and only two (rarely
three) fused cusps on each of the four multicusps.

DESCRIPTION.— Counts: Those of the holotype
given first (left and right sides), followed by ranges
for all specimens in parentheses: gill apertures 7,
7 (all); prebranchial slime pores 17, 16 (14-17);
branchial pores 7, 7 (6-8); trunk pores 67, 66
(60-67); cloacal pores 2, 3 (2-3); caudal pores
12, 12 (11-14); tail pores 14, 15 (14-16); total
slime pores 105, 104 (97-105). Cusps on anterior

multicusps 2, 2 (2, 3); posterior multicusps 2, 2
(2-3); anterior unicusps 15, 16 (13, 17); posterior
unicusps 12, 12(1 1-16); total cusps 63 (61-68).

Morphometry: Values in thousandths of TL
given first for the holotype, followed by ranges
for all specimens (left side): preocular length 53
(52-59); trunk length 560 (545-585); tail length
189 (181-213); maximum body depth including
finfold 89 (74-97), excluding finfold 84 (73-91);
body depth at cloaca 69 (61-80); tail depth 82
(82-89). Morphometric data (Table 1) and counts
(Tables 2-6) are compared with similar data for
other seven-gilled Eptatretus from the Pacific
Ocean.

Body deeper than wide, width at midbody
about 1.6 in depth. Tail broadly ovate, its depth
slightly greater than body depth. Ventral finfold
well developed, originating well behind PCD; a
broad, thin finfold from posterior margin of clo-
aca around tail and dorsally until about over
anterior margin of cloaca (Fig. 5-1). Dorsal pro-
file of head sloping to a very blunt, nearly straight-
across rostrum; width of nasopharyngeal orifice
about equal to length of third barbels. First pair
of barbels about 63% of length of third pair; sec-
ond pair about 75%.

Color notes were not taken at time of capture
(December 1965); all specimens are now a uni-
form yellowish color, no doubt a result of fading.
The eyespots are no longer discernible, but pre-
ocular measurements were recorded by the se-
nior author in November 1973. Removal of in-
tegument over the right eye of a 287-mm
specimen shows the eye to be round, about 2.5
mm in diameter, with a small triangular pupil
with its base dorsad and slanting forward at a
slight angle to horizontal axis of body. No sen-
sory canals are evident on any specimen.

Despite the faded condition of all specimens,
the branchial apertures and most slime pores have
whitish borders. Usually one pore, plus an oc-
casional extra one, occurs adjacent to each BA.
Two pores are near the opening of PCD on four
specimens; three have one pore, and one has
none near PCD (as is the usual condition on other

FIGURE 5. Tail shapes (to scale) and patterns of occurrence of the last four trunk pores and cloacal and tail slime pores of
four species of seven-gilled Eptatretus from the Pacific Ocean: 1-9 E. carlhubbsii (T = tan color, P = purplish color); 1 — Holotype,
SIO 68-473, 961 mm TL; 2-9 Paratypes: 2-SIO 68-473, 813 mm TL; 3-USNM 227440, 900 mm TL; 4-USNM 233742,
955 mm TL; 5-CAS 50705, 1064 mm TL; 6-BPBM 27848, 1 160 mm TL; 7-CAS 50706, 908 mm TL; 8-SIO 82-63, 1 125
mm TL; 9-BPBM 27851, 830 mm TL; 10-Holotype, E. laurahubbsi, SIO 65-643, 375 mm TL; 1 1— Holotype, E. strahani,
MNHN 1978-462, 520 mm TL; 12-E. cirrhatus, 655 mm TL.

262

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16

Eptatretus). Space between the last trunk pore
and first cloacal pore is about equal to length of
cloaca (Fig. 1). Two or three slime pores lie over
cloaca in a straight line and equally spaced with
caudal pores. Prebranchial pores in a fairly
straight line; occasionally the first one to three
slightly depressed.

There is great variation in the arrangement of
GP and afferent branchial arteries (ABA) with
respect to the DM and branching of VA (Figs.
3-2 and 3-3). The number of GP along DM are
far more variable than in the other three species,
ranging from two to five. Also, length of VA
varies notably; in six specimens VA averages 8.1%
(6.3-9.3%) of length of DM, but in one 240-mm
specimen the length of VA was 14.4% of DM,
with three ABA leading off the left side and two
off the right. This variation is in marked contrast
to the regular arrangement of the branchial ap-
paratus of the three other species discussed herein.

The eight specimens of E. laurahubbsi are un-
usual within genus Eptatretus in the appearance
of the multicusps, apparently having only two
fused cusps in each series. Indeed, it is often a
highly subjective decision as to whether any of
the anteriormost cusps are fused as multicusps.
One specimen appears to have three fused cusps
in each of the anterior series and two in the pos-
terior series. In two specimens it is questionable
as to whether two or three cusps are fused in the
posterior rows. In other Eptatretus known from
the southern hemisphere the usual configuration
is three fused cusps on each of the four multi-
cusps; all Eptatretus known from the North
American Pacific coast have three in the anterior
and two in the posterior row, which are distinct
and clearly seen even in juveniles. A juvenile
(188 mm) E. stoutii (Lockington 1878) clearly
shows hard, well-developed cusps with the pat-
tern 3/2. Also, a 100-mm specimen of E. poly-
trema (Girard 1855) from Chile has three dis-
tinctly fused cusps in each multicusp; adults of
this species attain a total length of at least 550
mm. Thus, the presence of only two fused cusps
in most specimens of E. laurahubbsi, and the
uncertainty regarding the number fused in the
others, is apparently not due to immaturity or
small size.

All of our study specimens are juveniles, but
it is highly probable that adults exceed one meter
in length. The longest (holotype), although 373
mm TL, contains minute eggs, seen with diffi-
culty under magnification. In another Eptatretus

(undescribed) from the Gulf of California, a spec-
imen of this size may have fully developed go-
nads and large eggs. Gumersindo Revuelta, a for-
mer student at the University of Chile,
Valparaiso, in an unpublished thesis (1976),
sketched and gave limited data on a very large
hagfish, slightly exceeding a meter in length (from
scale provided with sketch). He tentatively iden-
tified it with the giant Eptatretus taken at Wake
Island (from data sent to him by Hubbs), prob-
ably because of its gigantic size compared to oth-
er species from Chilean waters. Revuelta had at
least two females, both apparently very large, for
he sent to Hubbs (in 1976) two large eggs: one
68 by 16.5 mm from Valparaiso, and one 72 by
16.8 mm from "Juan Fernandez" (presumably
the island). We presume these large specimens
to be adults of E. laurahubbsi because Revuelta
reported the multicusps as 2/2, and his limited
data are in close agreement to those of our spec-
imens taken in the same vicinity. Also, in E.
carlhubbsi the ventral finfold is entirely absent,
but Revuelta's sketch shows a ventral finfold
originating a little behind the anterior third of
the body. In our juveniles a pronounced finfold
originates variably between the midpoint and an-
terior third of the body; the exact point of origin
is indeterminate because of wrinkling during
preservation.

ETYMOLOGY.— We dedicate this unique species
to our friend and and co-worker, Laura Clark
Hubbs, who contributed in so many ways to the
life and works of her husband, Carl Leavitt
Hubbs.

Eptatretus strahani new species

HOLOTYPE. -MNHN 1978-462, female, 520 mm TL, con-
taining eggs of about 4.5 mm in length, taken at 14°00'N,
120°1 8'.2"E, South China Sea near Lubang Island, Philippines,
in a trap net at 189 meters, Station 22 Musorstom Expedition,
21-22 March 1976, 1800-0600 hours.

PARATYPES (and remaining material examined; all taken with
the holotype). -MNHN 1981-722, female, 420 mm TL; SIO
81-116, female, 265 mm TL, male, 450 mm TL; USNM 227442,
male, 465 mm TL.

ADDITIONAL MATERIAL. -UPZM 1981-809, 400 mm TL;
UPZM 1981-811, 480 mm TL. Total lengths, comparisons,
and identifications were made by Prof. Reynaldo de La Paz,
University of the Philippines, Diliman, Quezon City, Philip-
pines, based on methods and data provided by us.

DIAGNOSIS.— A seven-gilled Eptatretus having
no eyespots, a well-developed ventral finfold, and
three fused cusps on each of the four multicusps.

DESCRIPTION.— Counts: Those of the holotype
given first, followed by ranges in parentheses for

MCMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES

263

all five specimens (both sides counted): gill ap-
ertures 7, 7 (all); prebranchial slime pores 14, 16
(13-16); branchial pores 6, 6 (6-7); trunk pores
45, 47 (45-48); cloacal pores 4, 3 (3-4); caudal
pores 7, 7 (6-8); total tail pores 11,11(10-12);
total slime pores 76, 79 (76-80). Cusps on mul-
ticusps 3, 3 (all); anterior unicusps 11,11 (9-1 1);
posterior unicusps 9, 9 (8-10); total cusps 52 (47-
52).

Morphometry: Values for holotype given first,
followed by ranges for all five specimens, in thou-
sandths of total length: preocular length (no eye-
spots); prebranchial length 23 1 (2 10-23 1); bran-
chial length 81 (69-83); trunk length 500 (500-
537); tail length 196 (174-202); body depth in-
cluding finfold 115(101-117); excluding finfold
95 (94-105); body depth over anterior margin of
cloaca 88 (77-94); tail depth 119 (109-125).
Morphometry (Table 1) and counts (Tables 2-
6) are compared with similar data for other sev-
en-gilled Eptatretus from the Pacific Ocean.

Body deeper than wide, deepest at midsection.
Ventral finfold well developed, extending from
about midbody to cloaca, its length about 31%
of TL. Tail margin quite thin posterior to cloaca,
extending around tail to dorsal surface, ending
at about a vertical from posterior end of cloaca.
Ventral outline of tail forms a nearly straight line,
ending with an abrupt curvature up and around
end to dorsal aspect. This shape is in marked
contrast to the gradual curvature of tails of the
other three species treated herein (Fig. 5).

Dorsal profile of head sloping steeply to snout;
rostrum more rounded than in E. carlhubbsi or
E. laurahubbsi; width of nasopharyngeal opening
about 60-80% of length of first pair of barbels.
First two pairs of barbels nearly equal in length;
respectively, about 66% and 72% of length of
third pair. First barbel, left side, of a 420-mm
female is bifurcate to near base, with posterior
branch shorter. Since we have occasionally seen
this bifurcation in other hagfishes (usually near
the tip, and always on only one barbel of the six),
we assume this form is the result of an injury
rather than some genetic malformation.

No eyespots are visible on any specimen (about
30 months after capture). Since the body color
is still fairly dark we have assumed that little or
no fading has occurred, and that the unpig-
mented eyespot area should still be visible if
present in life. No notes regarding eyespots were
made at time of capture. Removal of overlying
integument on holotype shows eye to be ovate

(3.4 by 2.4 mm) and slanted ventrodorsally at
about a 45° angle; pupil more rounded (1.4 by
1.1 mm).

Sensory canals are present in two groups on
each side of the head before and behind the area
where eyespots normally occur (Fig. 4). One group
of sensory canals is found near the bases of the
third pair of barbels, anterior to embedded eyes,
another group slightly posterior to eyes. Anterior
group consists of five more-or-less longitudinal
lines 1-3 mm long; canals of posterior group both
longitudinally and horizontally arranged, those
on top of head tending to be more horizontal.
No canals extend across dorso-medial line.

Color of holotype (in preservative) a light
brown, all paratypes a darker brown, the smallest
the darkest. No discernible whitish rings around
slime pores or GA on larger specimens, but the
GA of the smallest one has distinctly pale mar-
gins. Finfold anterior to cloaca is same color as
body, but tail has a very narrow, pale margin
extending a short distance forward on the dorsal
surface.

The line of the anterior prebranchial slime pores
is straighter in this species than in the other three
discussed; two specimens have only slight cur-
vature, and no anterior pores are markedly ele-
vated above adjacent ones in the prebranchial
series. Space between last trunk and first cloacal
pore about 65% of length of cloaca; spacing is
variable with degree of slant or elevation of first
cloacal pore (Fig. 5). Cloacal pores form a distinct
dorsoventral slant on left side of holotype, but
not on right; slanting is variable on paratypes.

Most GA are shaped as slits, slanting ventro-
dorsally; this shape could be an artifact of pres-
ervation, but the GA may be made to assume a
rounded form only by considerable pulling and
squeezing of surrounding flesh; all the slime pores
below GA are rounded.

Three to five GP lie anterior to tip of DM (Fig.
3); two to four lie between that tip and branching
of VA, and none posterior, although one GP of
the smallest specimen (265 mm TL) lies just at
the branching. Length of VA 6.4% (5.4-7.6%) of
DM length; DM length 26% (25-27%) of TL, its
width 15% (13-16%) of its length. Distance be-
tween tip of DM and branching of VA 9% (7.2-
10.9%) of DM length.

ETYMOLOGY.— We are pleased to dedicate this
new species to Ronald Strahan in acknowledg-
ment of his important contributions to the study
of Myxinidae.

264

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16

Eptatretus cirrhatus (Bloch and Schneider, 1801)

Petromyzon cirrhatus BLOCH AND SCHNEIDER, 1801:532 (orig-
inal description fide Forster ms 11:24 [habits: New Zealand]).

Homea banksii FLEMING, 1822:374 (South Seas [presump-
tive]).

Bdellostoma Forsteri MULLER, 1 834:7 U 80 (anatomy; char-
acters in key; reference to Petromyzon cirrhatus Bloch);
SCHNEIDER, 1 880: 1 1 5 (status uncertain; based on a poor
specimen).

Bdellostoma heptatrema MULLER, 1 834:7 (original description;
New Zealand).

Bdellostoma cirrahatum GUNTHER, 1870:511 (synonymy, in
part; diagnosis, in part; distribution [New Zealand only]);
HUTTON, 1872:87 (characters; color reddish brown, white
around mouth; common Australia and South Africa [mis-
identified with E. heptatrema]); PUTNAM, 1874:156 (in part;
New Zealand; 7 gill slits); SCHNEIDER, 1880:115 (in part;
doubts on status); ADAM AND STRAHAN, 1963:6 (6 or 7 pairs
of gills; average length 480 mm; South Pacific, common off
New Zealand).

Homea cirrhata GARMAN, 1899:341, 345, 349, 419 (synony-
my; nomenclature); DEAN, 1904:21 (in part; synonymy; New
Zealand).

Heptatrema cirrata [sic] HUTTON, 1904:55 (listed; New Zea-
land).

Eptatretus cirrahatus BERG, 1906:173 (in part; New Zealand);
WAITE, 1909:2 (description; behavior; average length 680
mm; Timaru and Chatham Islands; New Zealand); GRAHAM,
1965:67 (plentiful on North Otago Shelf, New Zealand; often
takes baited hooks); HEATH AND MORELAND, 1 967:30 (shore
to 1 800 ft; more abundant south of Hawke Bay than else-
where in New Zealand); WHITLEY, 1 968:4 (synonymy); SCOTT,
GLOVER, AND SOUTHCOTT, 1 974: 1 9 (New Zealand, New South
Wales, S.E. Australia); FERNHOLM, 1 974:35 1 (in shallow water,
New Zealand); FERNHOLM AND HOLMBERG, 1 975:253 (struc-
ture of eye, comparative; Kaikoura, S. Island, New Zealand);
STRAHAN, 1975:145 (key; description; ranges of counts and
body proportions).

Heptatretus banksii, REGAN, 1912:534, 536 (comparisons; syn-
onymy; diagnosis; D'Urville Is., Queen Charlotte Sound,
New Zealand.).

MATERIAL EXAMINED (counts and measurements both tak-
en).-SIO 81-105, two males, 488 and 655 mm TL, three
females 481-636 mm TL, 42°24'S, 1 73°4 1 'E, no data on depth
or date of capture, received from J. A. F. Garrick, Zoology
Department, Victoria University of Wellington, New Zealand,
1 Nov. 1972; SIO 62-482-4A, two females, 577 and 580 mm
TL, received from L. R. Richardson, Wellington, New Zealand,
25 March 1959, no data on depth of capture; ZIN 717-966,
male,595mmTL,40019'S, 172°15'E, 160- 172 meters, ISJan.
1965.

Counts only taken: AMS I 15527-001, three males, 254-
452 mm TL, 26°32'S, 153°51'E, agassiz trawl, 175 fms (320
m), 27 July 1968; AMS Kapala Station 71-07-03, female, 505
mm TL, 33°33'-37'S, 152°01'-151°57'E, 205 fms (375 m), 21
April 1971; AMS Kapala Station 71-08-05, male, 265 mm TL,
female, 552 mm TL, 33°11'S, 152°23'E, otter trawl, 310 fms
(567 m), 29 April 1971; AMS Kapala Station 71-1 1-07, two
females, 49 1 , 5 1 4 mm TL, 34°40'-35°0 1'S, 1 5 1 ° 1 0'-07'E, otter
trawl, 300 fms (549 m), 7 July 1971; AMS Kapala Station 71-
1 1-08, female, 410 mm TL, 34°56'-35°02'S, 151°06'-05'E, ot-
ter trawl, 160 fms (194 m), 8 July 1971; AMS Kapala Station

71-1 1-10, male, 447 mm TL, 35°1 l'-37'S, 150°45'-42'E, otter
trawl, 230 fms (421 m), 8 July 1971; AMS Kapala Station 71-
12-04, female, 546 mm TL, 33041'-49'S, 151°53'-47'E, otter
trawl, 245-250 fms (448-457 m), 20 July 1971; AMS Kapala
Station 71-12A-06, three females, 374-503 mm TL, 35°25'-
29'S, 150°50'-48'E, otter trawl, 300 fms (549 m), 2 Aug. 1971.

DIAGNOSIS.— A seven-gilled Eptatretus having
a vestigial ventral finfold, small but prominent
eyespots, white around the mouth, pale rings
around branchial apertures and slime pores; three
fused cusps on each multicusp; sensory canals
may occur.

DESCRIPTION.— Despite its being the first hag-
fish described from the Pacific Ocean, the liter-
ature contains minimal data on morphology and
counts. Waite (1909) and Graham (1965) pro-
vided descriptions and accounts of behavior, but
no meristic data. Strahan (1975) listed only ranges
of counts and percentages of total length for cer-
tain body measurements for 13 specimens. We
offer morphometry based on only the eight spec-
imens available to us, but include counts on 22
specimens examined by Carl L. Hubbs in 1971
at the Australian Museum, Sydney.

Counts: Averages followed by ranges in pa-
rentheses, both sides counted: gill apertures 7
(all); prebranchial slime pores 17-18 (16-20);
branchial pores 7 (6-8); trunk pores 48-49 (46-
53); cloacal pores 3 (2-5); caudal pores 9 (6-1 1);
total tail pores 12 (10-14); total slime pores 86
(83-90). Three fused cusps on each of the four
multicusps; anterior unicusps 9 (8-1 1), posterior
unicusps 8 (7-9); total cusps 46 (43-51).

Morphometry: Averages followed by ranges in
parentheses, in thousandths of TL, for eight spec-
imens: preocular length 60 (52-67); prebranchial
length 225 (214-239); branchial length 76 (69-
89); trunk length 550 (525-563); tail length 154
(135-168); maximum body depth including fin-
fold 93 (84-102); excluding finfold 91 (81-102);
body depth over cloaca 67 (57-74); tail depth 82
(77-9 1); body width at mid-prebranchial section
50 (46-55).

Dorsal profile of head sloping gently down-
ward, face sloping at about a 45° angle; naso-
pharyngeal opening about equal to or slightly less
than length of first barbel. Average length of first
pair of barbels about 69% and second pair about
75% of the length of the third pair. Body rounded
anteriorly, becoming more laterally compressed
and deeper posteriorly; tail bluntly rounded,
spatulate, its depth slightly less than greatest depth

MCMILLAN AND WISNER: NEW SPECIES OF PACIFIC HAGFISHES

265

of body (Table 1). Ventral finfold narrow and
relatively short, its length about 30% of TL, ex-
tending from well behind midbody to cloaca.

Color of our specimens in preservative varies
from light chocolate to dark brown; the most
lightly colored one is strongly mottled anteriorly
with small, irregular pale spots and patches. Waite
(1909) stated that "the colour varies from blue
to bluish violet [presumably fresh material]. Some
examples show irregular white spots and mark-
ings; the ventral finfold and the margin of the
tail may also be white." On our material, pre-
served at least ten years prior to our examination,
the poorly developed finfold has a narrow pale
margin on two specimens, but not on the other
six; color is highly variable. Tail of one specimen
is irregularly margined with pale areas; pale rings
around most of the gill apertures and slime pores.

Small but plainly visible eyespots are present
on all specimens. Removal of integument cov-
ering right eyespot of a specimen 577 mm TL
reveals the embedded eye as elliptical (6.3 by 3.2
mm) and slanting forward ventrodorsally at about
a 45° angle; pupil small, more rounded (about
1.9 by 1.4 mm).

Sensory canals, only as posterior series (in the
specimens available to us), are located close be-
hind eyespots (Fig. 4). Canals are readily iden-
tifiable on only three of eight specimens (488-
655 mm TL) and only on the left side of largest
one; they are similar in size, form, and arrange-
ment to those of E. strahani. On two specimens
(580 and 597 mm TL) positive identification of
canals is prevented by presence of much scarring.

The anterior prebranchial slime pores on all
specimens form a downsloping curve; a high in-
cidence of irregular spacing of these pores occurs.
On left side of four specimens, first or third pore
is notably elevated or very closely spaced. On
one specimen four pores form an almost-square
pattern; all apparently lead from a single slime
gland. In most Eptatretus from the Pacific Ocean,
the usual number of branchial slime pores is one
less than the number of GA, since ordinarily
there is no pore associated with the PCD. In E.
cirrhatus there is a high incidence of extra pores
and irregular spacing of slime glands in the bran-
chial region, especially near the PCD.

Posterior two or three GA, left side, curve gently
downward on seven of eight specimens, but only
on the right side of one. On a 480-mm juvenile
female the normally confluent openings of the

seventh GA and PCD are distinctly separate; such
separation also occurs infrequently in other
species of Eptatretus. Usually five, occasionally
four, GP lie anterior to tip of DM. Length of DM
27% of TL, its width 13% (12-14%) of its length;
somewhat flattened posteriorly, depth is 80% (75-
88%) of its width. The shape of the tip of the
DM varies somewhat from that of the other three
species discussed herein (Fig. 3). Distance be-
tween tip of DM and branching of VA about
7.4% (4.8-10.8%) of DM length; VA usually
greater in width than in length, in contrast to that
of the other three species. No ABA lead ofFbefore
branching of VA in the eight specimens available
to us.

A 655-mm TL female contains about 50 large
eggs, ranging from 29 to 32 mm long by about
10 mm in diameter; this is an unusually large
number of maturing eggs. All are still in the mes-
entery which is attached to the body wall; ter-
minal anchor filaments and hooks are not present
on any egg.

ACKNOWLEDGMENTS

We are deeply grateful to the late Carl L. Hubbs
for his efforts in amassing much of the material
on hagfishes, without which this study might
never have been accomplished. Also, we are most
grateful to the following persons and agencies:
Jacques Forest, Museum National d'Histoire
Naturelle, Paris, and Professor Reynaldo M. de
La Paz, University of the Philippines, Diliman,
Quezon City, for material of E. strahani; J. A.
F. Garrick, Victoria University, Wellington, New
Zealand, and L. R. Richardson, Wellington, for
specimens of E. cirrhatus; and Richard H. Ro-
senblatt, for the holotype and one paratype of E.
carlhubbsi. Other material of E. carlhubbsi was
provided by Steven Kramer, National Marine
Fisheries Service, Hawaii; Robert Moffitt, NMFS
Field Station, Mangilao, Guam; Paul Strushaker,
Honolulu; and John E. Randall and Arnold Su-
zumoto, Bernice P. Bishop Museum, Honolulu.

All specimens ofE. laurahubbsi were taken by
the junior author on Cruise 1 2, R/V Anton Bruun,
Southeastern Pacific Biological Oceanography
Program, sponsored by the National Science
Foundation. The holotype and one paratype of
E. carlhubbsi were taken on the Scripps Styx ex-
pedition, sponsored by the National Science
Foundation through a grant (GB-7596) to R. H.
Rosenblatt and W. A. Newman. We greatly ap-

266

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 16

predate the efforts of all persons involved. R. H.
Rosenblatt critically read the manuscript.

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CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, California 94 1 1 8

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 17, pp. 269-282, 5 figs.

December 11, 1984

NEW AND NOTEWORTHY ADDITIONS TO THE
MELASTOMATACEAE OF PANAMA

By
Frank Almeda

Department of Botany, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118

ABSTRACT: Descriptions, discussions, diagnostic illustrations, and distributional notes are provided for
five new species of Melastomataceae (Clidemia tetrapetala, C. trichosantha, Graffenrieda bella, Miconia
arboricola, and Topobea calophylla). Based on a study of the first known flowering collections, Topobea
elliptica is transferred to Blakea and provided with an amplified description. Notes are also presented on the
size and delimitation of these genera with special reference to Panama.

INTRODUCTION

The first and only floristic treatment of the
Panamanian Melastomataceae was published
some 25 years ago (Gleason 1958). The manu-
script for that study was completed several years
before it appeared in print. Gleason's study is a
very useful account that unfortunately suffers
from a lack of adequate field study and a paucity
of local material for many of the species attrib-
uted to Panama. The many new collections of
Melastomataceae generated by the heightened
botanical exploration of Panama during recent
years has underscored the need for an updated
supplement. Such an undertaking is still pre-
mature in view of the many undescribed taxa
represented by collections that are too incom-
plete to serve as a basis for formal descriptions.

In this interim report I propose new species in
Clidemia, Graffenrieda, Miconia, and Topobea,
present a new combination in Blakea, and pro-
vide brief notes on the size and delimitation of
these genera in Panama.

Blakea P. Browne

Blakea and the closely related Topobea con-
stitute the tribe Blakeae, which is characterized
by baccate fruits, ovoid to pyramidal seeds, and
axillary, 6-merous flowers that are individually
subtended by two pairs of decussate bracts in-
serted at the base of the hypanthium. These gen-
era exhibit many parallel variations in foliar
characters, floral bract morphology, and calyx
development. The traditional distinction be-
tween Blakea and Topobea is based on the an-
droecial characters summarized in the following
couplet:

Anthers short, oval, oblong, or elliptic, blunt
or broadly rounded at the summit with two
typically well-separated (often minute) api-
cal pores Blakea

Anthers linear-oblong to lanceolate or subu-
late, the dorsally inclined apical pores ap-
proximate and often confluent at anthesis
Topobea

[269]

270

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17

In addition to the three species of Blakea de-
scribed by Almeda (1974, 1980) and the generic
transfer presented below, Blakea tuberculata
Donn.-Smith of Costa Rica has recently been
collected in western and central Panama (Chi-
riqui: Folsom et al. 4838, CAS; Code: Knapp
5334, CAS; Panama: Sytsma 1154, CAS). These
additions increase the number of described Pan-
amanian species to thirteen.

Blakea elliptica (Gleason) Almeda, comb. nov.

Topobea elliptica GLEASON, Phytologia 3:353. 1950.

TYPE.— Panama. Bocas del Toro, Robalo Trail, northern
slopes of Cerro Horqueta, elevation 6000-7000 ft [1828-2133
m], 5-7 Aug. 1947, Allen 5001 (holotype MO!; photo CAS!).
The type collection of this species consists of a branch with
four leaves and four more or less mature berries (one of which
is detached). When Gleason described this species from in-
complete material he was impressed by its sessile, clasping
leaves, which are reminiscent of those of Topobea brenesii
Standley, a rare Costa Rican endemic. Two recent flowering
collections from near the type locality in western Panama are
an exact vegetative match for the species that Gleason placed
in Topobea. The uniform, bluntly ovoid anthers of these new
collections are characteristic of Blakea, thus necessitating the
generic transfer proposed herein. Because all available descrip-
tions of this taxon are based on the fragmentary type collection,
an amplified description is presented below to facilitate rec-
ognition of this little-known cloud forest epiphyte.

Epiphytic shrub reportedly 2-4 m tall. Cauline
internodes glabrate, terete, becoming striate to
furrowed or cracked in age. Distal branchlets and
floral pedicels sparsely covered with a deciduous
indument of slender, spreading, glandular tri-
chomes and stoutly conic trichomes, the latter
typically copious on distal nodes and vegetative
buds. Mature leaves sessile, blades chartaceous
to coriaceous, entire, somewhat revolute on
drying, 5-1 1.5 cm long and 3-7.5 cm wide, el-
liptic-ovate to obovate, obtuse to rounded api-
cally and auriculate to cordate-clasping basally,
3-nerved with an additional but less conspicuous
submarginal pair, essentially glabrous through-
out or with a sparse scattering of conic and glan-
dular trichomes at the base of the blade below.
Flowers 6-merous, solitary or paired in the axils
of distal branches, pedicels 2.5-4.3 cm long. Flo-
ral bracts foliaceous, sparsely covered with a
mixture of blunt conic and spreading glandular
trichomes on the inner and outer surfaces; outer
bracts 3-5 -nerved, essentially free to the base,
12-14 mm long and 13-15 mm wide, broadly
elliptic-ovate to suborbicular, obtuse to rounded
or somewhat retuse apically; inner bracts (at an-

thesis) connate basally for about 4-5 mm to form
a collar that tightly envelops the ovary, the free
lobes ± semicircular, 7 mm long and 12-13 mm
wide basally between sinuses. Hypanthia (at an-
thesis) glabrous, campanulate, 8-1 1 mm long to
the torus. Calyx lobes triangular, ± entire, most-
ly recurved, 5-6 mm long and 5 mm wide ba-
sally, apically beset with a mixture of spreading
glandular and sessile globular trichomes. Petals
thin and ± translucent on drying, mostly gla-
brous with a moderate but somewhat concen-
trated abaxial patch of brown verrucose excres-
cences, reportedly white with pink blotches,
elliptic-ovate to obovate and rounded apically,
entire to irregularly ciliolate, 20-23 mm long and
14-18 mm wide. Stamens isomorphic, free and
declined to one side of the flower, filaments gla-
brous, 4.5-6 mm long and 1 .5 mm wide; anthers
yellow, ovoid, laterally compressed, biporose and
rounded apically, 5 mm long and 1.5-2 mm wide,
connective thickened dorsally and dilated basally
about 0.5 mm above point of filament insertion
into a short deflexed truncate appendage about
0.5-1 mm long. Style straight, glabrous, con-
spicuously exserted, 11-14 mm long; stigma
truncate. Berry 13-15 mm long to the torus and
14-15 mm in diameter, glabrous and subglobose.
Seeds beige, cuneate to clavate, or narrowly pyr-
iform, mostly 1 mm long.

ADDITIONAL SPECIMENS EXAMINED.— Panama. Chiriqui: Bo-
cas del Toro border along Continental Divide NE of Cerro
Pate Macho, above Palo Alto in windswept elfin forest. 8°47'N,
82°2 1 ' W, 2200 m, 1 5 Mar. 1 982, Knapp et al. 4233 (MO); end
of road past Palo Alto NE of Boquete in forest along ridge;
elev. 6200-6800 ft [1890-2073 m], 8 Feb. 1979, Hammel6048
(MO).

DISTRIBUTION. — Western Panama. Known
only from the northern slopes of Cerro Horqueta
and the region NE of Cerro Pate Macho above
Palo Alto at elevations of about 1800-2100 m.

This species is apparently rare and localized,
as evidenced by the few specimens collected over
the past three decades. It is among the most dis-
tinctive of the Central American species by vir-
tue of its sessile, clasping leaves, and deciduous
indument of spreading glandular and blunt conic
trichomes on distal branchlets, pedicels, floral
bracts, and vegetative buds. The petals of this
species are remarkably thin and translucent when
dry and have a peculiar abaxial cluster of brown
verrucose excrescences. Additional collections
and field observations will be needed to deter-

ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA

271

mine whether this petal feature is a consistent
diagnostic character or an artifact of pressing and
drying.

Clidemia D. Don

Clidemia is one of approximately 30 genera
constituting the tribe Miconieae. The generic
classification of this complex tribe is difficult be-
cause it contains species groups that intergrade
across generic lines that have been defined on
the basis of one or few characters. The evolu-
tionary development of characters used in ge-
neric delimitation of the Miconieae is unclear,
and some of the genera as currently delimited
are probably polyphyletic. Until a reassessment
of generic limits can be accomplished, Clidemia
is defined in the traditional sense to include those
berry-fruited species with axillary inflorescences,
more or less obtuse petals, linear to subulate an-
thers, wingless hypanthia, and flowers that are
not individually subtended by two pairs of de-
cussate bracts.

Of the approximately 165 species of Clidemia
recorded for tropical America, 30 are now known
from Panama. This count includes the two new
taxa proposed below.

Clidemia tetrapetala Almeda, sp. nov.

(Figure 1)

Frutex 1-2 m. Ramuli teretes sicut petioli in-
florescentiafoliorum subtus venae primariae pilis
stipitato-stellatis densiuscule ornati et pilis stel-
latis sessilibus modice vel dense induti. Petioli
10-27 mm longi; lamina 5-9.5 x 2.1-3.9 cm
ovato-elliptica apice gradatim acuminato basi
rotundata vel subcordata, chartacea et Integra,
5-7-plinervata. Inflorescentiae pauciflorae info-
liorum superiorum axillis oppositis, axe plerum-
que 1.5-2.5 cm longo; flores 4-meri plerumque
subsessiles (pedicellis supra articulationem 0.5
mm longis), bracteolis 1-1.5 mm longis lineari-
bus. Hypanthium (ad torum) 1.5 mm longum
dense stellato-puberulum et pilis laevibus glan-
duliferis patentibus 0.5-1 mm longis modice in-
termixtus; lobis 1 mm longis triangularibus, den-
tibus exterioribus subulatis 3 mm eminentibus.
Petala 5-6 x 1.5-2 mm oblonga glabra (apice
late obtuso). Stamina isomorphica glabra; fila-
menta 2 mm longa; anther arum thecae 1.5 x
0.5 mm oblongae, poro dorsaliter inclinato; con-
nectivum nee prolongatum nee appendiculatum.

Stylus 5.5 mm glaber; stigma truncatum; semina
0.5-1 mm pyriformia.

Shrubs 1-2 m tall. Older cauline internodes
glabrate and terete. Distal branchlets, vegetative
buds, juvenile leaves, and inflorescences copi-
ously covered with sessile, and stipitate-stellate
trichomes, the distal cauline internodes sparingly
intermixed with spreading glandular hairs. Leaves
of a pair slightly unequal in size, otherwise iden-
tical in shape and vestiture. Leaf blades flat to
somewhat bullate above when dry, chartaceous,
entire, 5-9.5 cm long and 2. 1-3.9 cm wide, ovate-
elliptic, acuminate apically and rounded to sub-
cordate basally, 5-7-plinerved with a network of
secondary and tertiary nerves below, sparsely pu-
bescent above with a mixture of appressed sim-
ple trichomes and spreading stalked stellate tri-
chomes, lower leaf surfaces copiously stellate
pubescent on the elevated primary nerves but
with a moderate to sparse mixture of sessile and
stalked stellate trichomes on and between the
secondaries; petioles 10-27 mm long and 1-1.5
mm broad. Inflorescence axillary, typically a lax
few-flowered modified dichasium 1.5-2.5 cm
long; bracteoles sessile, 1-1.5 mm long and most-
ly less than 0.5 mm wide, linear-lanceolate to
linear-oblong, mucronate apically, glabrous above
and sparsely stellate pubescent below. Pedicels
terete, mostly less than 0.5 mm long and disar-
ticulating at the tribracteolate node closely sub-
tending each flower. Hypanthia (at anthesis) nar-
rowly campanulate, 1.5 mm long to the torus,
copiously covered with a mixture of stellate and
spreading glandular trichomes. Calyx lobes (on
fruiting hypanthia) widely spreading, persistent,
triangular, entire, 1 mm long and 1 mm wide
basally; exterior calyx teeth subulate, mostly 3
mm long, stellate pubescent and completely ob-
scuring external surface of the calyx lobes. Petals
4, glabrous, reportedly white, linear-oblong,
broadly obtuse apically, entire, 5-6 mm long,
1.5-2 mm wide. Stamens 8, isomorphic, erect at
anthesis; filaments glabrous, ligulate but abruptly
constricted distally, 2 mm long; anthers 1.5 mm
long and 0.5 mm wide, yellow when dry, linear-
oblong, truncate to rounded distally with a cir-
cular dorsally inclined apical pore; connective
simple, somewhat thickened dorsally but not di-
lated or prolonged below the thecae. Ovary to-
tally inferior, 4-celled, essentially glabrous at the
summit but sparsely setose surrounding the sty-
lar scar. Style 5.5 mm long; stigma truncate. Ber-

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17

FIGURE 1 . Clidemia tetrapetala Almeda. A, habit, x ca. %; B, representative leaf (lower surface), x ca. %; C, mature berry, x
ca. 4; D, petal, x ca. 6; E, stamens, 3A lateral view (left) and dorsal view (right), x9; F, seeds, x ca. 7. (A from Folsom 4871;
B-F from Folsom 6106.)

ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA

273

FIGURE 2. Clidemia trichosantha Almeda. A, habit, x ca. %; B, representative leaf (lower surface), x ca. 3/t; C, representative
flower, buds, bracts and bracteoles, x ca. 4; D, petal, x 7; E, stamens, lateral view (left) and ventral view (right), x 9; F, seeds,
x ca. 14. (A-E from the holotype; Ffrom Hammel 965.)

ry reportedly Opaque green but turning translu- TYPE.— Panama. Bocas del Toro/Chiriqui Border: Cerro

Cent red at maturity, globose, 5-6 mm long tO Colorado, along intersection of Bocas Road with main ridge

the torus, 5-6 mm in diameter. Seeds narrowly Load' "-8 km "^S^rKTJS1?,"0"!,^

J Toro, elevation 1400-1700 m, 24 Oct. 1977, Folsom 6106

pyriform, brown, smooth, vernicose, and mostly (holotype: CAS!; isotype: MO).

0.5—1.0 mm long. ADDITIONAL SPECIMENS EXAMINED.— Panama. Bocas del

274

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17

Toro/Chiriqui Border: Cerro Colorado, 1 1.2 km along ridge
road from main road to Escopeta, 1700 m, 16 Aug. 1977,
Folsom 4871 (CAS, MO). Chiriqui: Cerro Colorado on Con-
tinental Divide, 1400 m, 25 Jul. 1979, Antonio 1416 (CAS).

DISTRIBUTION.— Known only from Cerro Col-
orado in western Panama at elevations of 1400-
1700 m. All cited specimens are in flower; only
the type has mature fruits.

Among Panamanian species of Clidemia with
4-merous flowers, C. tetrapetala is distinguished
by its lance-ovate, 5-7-plinerved leaves, linear-
oblong petals, and mixture of spreading glan-
dular and stellate hypanthial trichomes. The in-
florescence of this species is also notable for its
elongate distal peduncles that can readily be mis-
taken for floral pedicels. The actual pedicels in
this species are very short and disarticulate at
tribracteolate nodes that closely subtend the
flowers.

On the basis of its 4-merous flowers, well de-
veloped calyx teeth, and equal to subequal prin-
cipal leaves in each pair, this species is provi-
sionally placed in section Sagraea as defined by
Cogniaux (1891). Among species of this section,
C. tetrapetala bears a superficial resemblance to
C. saltuensis Wurdack of Venezuela which is re-
portedly known only from the type (Wurdack
1973). The latter species differs most conspicu-
ously in having ovate-elliptic, basally obtuse
leaves, stellate-pinoid hypanthial trichomes,
shorter (0.7 mm), erect calyx teeth, narrowly ob-
long anther thecae, and abaxially mucronate pet-
als.

Clidemia trichosantha Almeda, sp. nov.

(Figure 2)

Frutex 1-2.5 m. Ramuli teretes primum sicut
petioli laminae subtus inflorescentia hypan-
thiaque dense setosi pilis l-3(-4) mm longis de-
mum glabrati. Folia in quoque pari in dimen-
sionibus disparilia (2:1); petioli 0.2-2.2 cm longi;
lamina 3-14.3 x 1.5-5.8 cm elliptica apice acu-
minata basi interdum obtuse vel rotundata vel
paulo asymmetrica, chartacea et denticulata, 5-
7-plinervata, supra sparse vel modice strigosa.
Inflorescentiae ca. 1.5-3 cm longae infoliorum
superiorum axillis; flores 5-meri, pedicellis 1-1.5
mm longis, bracteolis 1.5-3 mm longis lanceo-
latis amplexicaulibus. Hypanthium (ad torum)
2-2.5 mm longum; lobis interioribus late deltoi-
deis, dentibus exterioribus setosis 1-2 mm emi-
nentibus. Petala glabra 4-4.5 x 7.5 mm oblon-

go-lanceata. Stamina isomorphica glabra;
filamenta 2.5 mm longa; antherarum thecae
1.5 x 0.75 mm oblongae, connectivum nee pro-
longatum nee appendiculatum. Stylus 4.5 mm
glaber; stigma punctiforme; semina 0.5 mm cu-
neata.

Slender shrubs 1-2.5 m tall. Internodes terete,
glabrate at maturity, but vegetative buds, young
leaves, and distal branchlets moderately to
densely covered with smooth, eglandular,
spreading trichomes mostly l-3(-4) mm long.
Leaves of a pair usually unequal in size, other-
wise identical in shape and vestiture. Leaf blades
chartaceous, denticulate, elliptic, acuminate api-
cally, acute varying to obtuse, rounded or con-
spicuously oblique basally, 5-7-plinerved with a
network of secondary and tertiary nerves, the
innermost pair of primary nerves diverging from
the median nerve (0.6-)1.0-1.6 cm above the
petiole/laminar junction, moderately to sparsely
strigose or hirtellous above and moderately to
copiously hirsute below. Larger leaf at each node
5.8-14.3 cm long and 2.5-5.8 cm wide with pet-
ioles mostly 0.5-2.2 cm long. Smaller leaf 3-9.5
cm long and 1.5-4. 1 cm wide with petioles most-
ly 2-4 mm long. Inflorescence an axillary, mul-
tiflowered, modified dichasium with flowers
borne in pedunculate terminal glomerules; rachis
terete, 1.5-3 cm long, moderately to sparsely hir-
sute; bracts and bracteoles sessile, paired and
often fused basally into an amplexicaul nodal
collar, 1.5-3 mm long and 0.5-1.5 mm wide,
lanceolate to naviculiform, glabrous but irregu-
larly fringed with spreading setose trichomes.
Pedicels hirsute, 1-1.5 mm long. Hypanthia (at
anthesis) ± ovoid, 2-2.5 mm long to the torus,
copiously covered with smooth spreading tri-
chomes. Calyx lobes (on fruiting hypanthia) in-
conspicuous, erect, persistent, broadly deltoid,
entire, 1 mm long and 1.5 mm wide basally be-
tween sinuses; exterior calyx teeth setiform, 1-2
mm long, sparsely hirsute. Petals 5, glabrous,
reportedly translucent white, elliptic-lanceolate,
acute apically, entire, 4-4.5 mm long, 1.5 mm
wide. Stamens 10, isomorphic, apparently erect
at anthesis; filaments glabrous, ± linear-oblong
but constricted distally, 2.5 mm long; anthers 1.5
mm long and 0.75 mm wide, linear-oblong, trun-
cate distally; connective simple, not dilated or
prolonged below the thecae. Ovary ca. % inferior,
5 -celled. Style 4.5 mm long; stigma punctiform.
Berry purple at maturity, globose, 4-6 mm long

ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA

275

and 4-5.5 mm in diameter. Seeds cuneate, ±
smooth with verruculose angles and a promi-
nent ± translucent lateral raphe, 0.5 mm long.

TYPE.— Panama. Code: sawmill above El Cope, in forest
along stream east of sawmill on the Atlantic drainage, elevation
2300 ft (70 1 m), 27 July 1978, Hammel 4133 (holotype, CAS!;
isotype, MO).

ADDITIONAL SPECIMENS EXAMINED. — Panama. Cocl6: above
El Petroso sawmill at Continental Divide, N of El Cope, 1 3
May 1981, Sytsma & Andersson 4624 (CAS); area surrounding
Rivera sawmill, Alto Calvario, 7 km N of El Cope at Conti-
nental Divide, 25 Nov. 1977, Folsom & Collins 6473 (CAS);
near Continental Divide along lumbering road 8.4 km above
El Cope, 19 Jan. 1978, Hammel 965 (CAS); 7 km N of El
Cope around Rivera sawmill, 2 1 Dec. 1 977, Folsom et al. 7093
(CAS); El Cope on Pacific side about '/z hour walk from the
sawmill, 16 Oct. 1979, Antonio 2116 (CAS). Panama: forested
slope along El Llano-Carti road, 12 km from Panamerican
Highway, 10 Sept. 1976, D'Arcy 10617 (CAS). San Bias: El
Llano-Carti road, 12 mi from Panamerican Highway, 10 May
1981, Sytsma & Andersson 4493 (CAS). Veraguas: Cerro Tute,
30 Nov. 1979, Antonio 2928 (CAS).

DISTRIBUTION.— Presently known from wet
forests and stream margins at elevations from
700-1300 meters in central Panama. Flowering
specimens have been gathered in January, May,
July, and September.

The epithet for this species draws attention to
its most remarkable feature— the dense covering
of spreading, lustrous trichomes on the flowers
and peduncles of the inflorescence. The tri-
chomes, which are often basally flattened when
dry, are so copious that it is difficult to examine
the structure and posture of bracteoles and other
floral parts without a dissection of hydrated ma-
terial. Other characters that readily separate C.
trichosantha from its congeners include the mod-
ified dichasial inflorescence of pedunculate ter-
minal glomerules, the distally constricted fila-
ments, the cuneate seeds with verruculose angles,
and the sessile, paired bracts and bracteoles that
are often fused into amplexicaul collars.

A fruiting collection from Cerro Habu, San
Bias (Sytsma et al. 2673, CAS) appears to be a
variant or closely related, undescribed taxon but
its essentially glabrous leaves, glabrate berries,
and short (0.5 mm) calyx teeth fall outside the
range of variation here attributed to C. tricho-
santha.

The 5-merous, short-pedicellate flowers and
anisophylly of the new species dictate placement
into Cogniaux's section Calophysoides. Clidemia
trichosantha is unlike any of the species currently
placed in that section. I am, therefore, reluctant

to include it with an assemblage that may prove
to be artificial and can only suggest that no close
relatives are apparent. The combination of
smooth, spreading trichomes and elongate calyx
teeth is vaguely suggestive of the copiously hir-
sute variants of C. petiolaris (Schlecht. & Cham.)
Schlecht. ex Triana which, otherwise, differ
markedly in their lax, divaricately branched in-
florescence, subulate, apically notched anthers,
and pyriform, papillate seeds.

Graffenrieda DC.

Graffenrieda, a genus of about 40 species, is
one of five closely related merianioid genera oc-
curring in Panama. Gleason (1958) reported one
species (under the generic synonym Calyptrella
Naud.) in his treatment of Panamanian Melas-
tomataceae. The escalated botanical exploration
of Panama in the past two decades has increased
this total to about eight. In addition to the species
proposed below, G. galeottii (Naud.) L. Wms.,
G. gracilis (Triana) L. Wms., and G. micrantha
(Gleason) L. Wms. are known from Panama.
Collections of about four additional entities
probably represent undescribed taxa or range ex-
tensions of South American species. These re-
main unnamed because of incomplete material.

In the field, the superficial resemblance of Pan-
amanian species of Graffenrieda to Miconia is
striking. Graffenrieda is readily distinguished by
the combination of capsular fruits, calyptrate ca-
lyx, dorsally spurred, arcuate anthers, and linear,
pyramidate seeds.

Graffenrieda bella Almeda, sp. nov.

(Figure 3)

Arbor parva 5 m. Ramuli glabri et teretes. Pe-
tioli 0.5-1. 6 cm; lamina 4.7-8 x 2.1-4 cm ellip-
tica apice breviter subabrupteque acuminato basi
acuta vel cuneata vel biauriculata, supra glabra
vel sparse lepidota, subtus modice lepidota,
3(-5)-nervata, coriacea et Integra. Panicula 5-7
cm longa multiflora; flores 5(-6)-meri subsessiles
(vel pedicel Us 0.5-1 mm longis) ad ramulorum
apices glomerati (floribus ca. 3-6 in quoque glo-
merulo), bracteolis 0.5-1 mm longis subulato-
setosis evidenter mox caducis. Hypanthium (ad
torum) 3 mm longum; calyx in alabastris ma-
turis clausus 3-3.5 mm longus ad anthesim ir-
regulariter supra torum dehiscens. Petala 7-9 x
4-6 mm elliptico-ovata glabra. Stamina isomor-
phica glabra; filamenta 2.5-3 mm longa; an-

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17

FIGURE 3. Graffenrieda bella Almeda. A, habit, x%; B, representative leaves, lower surface (left) and upper surface (right),
x ca. %; C, mature hypanthia, x ca. 4; D, calyptrate calyx, x ca. 8; E, petal, x ca. 5; F, stamen (lateral view), x ca. 10; G,
seeds, x ca. 14. (A-B, D-Ffrom the holotype; C& G from Mori et al. 7581.)

therarum thecae 3-4.5 x 0.5 mm oblongo-
subulatae; connectivum vix (0.25 mm) prolon-
gatum, dente dorsali 0.5 mm longo acuto. Stylus
7.5-9.5 mm glaber; stigma punct iforme; semina
numerosa 1 mm longa, recta paulo angulata.

Trees to 5 m tall. Cauline internodes glabrous
and terete; the nodes on defoliated branchlets

somewhat swollen with age. Leaf blades coria-
ceous, entire, 4.7-8 cm long and 2. 1-4 cm wide,
broadly elliptic, apically acuminate basally acute
to cuneate or biauriculate with well-developed
reflexed marginal lobes ca. 2 mm above the pet-
iole-laminar junction, glabrous to sparsely lepi-
dote above and moderately lepidote below at

ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA

277

maturity, uniformly 3 -nerved (excluding the in-
conspicuous submarginal pair); petioles 0.5-1.6
cm long and 1-1.5 mm broad. Inflorescence a
multiflowered panicle mostly 5-7 cm long (in-
cluding the peduncle), the uppermost units com-
monly glomerulate; bracteoles sessile, narrowly
subulate to acicular, 0.5-1 mm long, apparently
early deciduous and mostly absent at anthesis.
Pedicels 0.5-1 mm long. Hypanthia (at anthesis)
narrowly campanulate to subcylindric, 3 mm long
to the torus. Calyx acute, apiculate and closed in
bud, 3-3.5 mm long rupturing near the torus and
falling away as a unit at anthesis. Calyx teeth (on
mature hypanthia) evident as blunt tuberculi-
form projections mostly 0.25-0.5 mm long. Pet-
als mostly 5, but varying to 6 in some flowers,
glabrous, white, elliptic to ovate, entire, acute
apically, cuneate to clawed basally, 7-9 mm long
and 4-6 mm wide. Stamens mostly 10 but some-
times 12, isomorphic; filaments glabrous, ligu-
late, 2.5-3 mm long and 0.5 mm wide; anthers
3-4.5 mm long and 0.5 mm wide, yellow, sub-
ulate, the apical pore ± truncate; connective pro-
longed ca. 0.25 mm below the thecae and mod-
ified dorsally (at filament insertion) into an acute
spur mostly 0.5 mm long. Ovary '/4 inferior,
3-celled, ovoid, glabrate but copiously lepidote
distally. Style straight to somewhat curved,
glabrous, 7.5-9.5 mm long; stigma punctiform.
Hypanthia (at maturity) prominently costate, 5
mm long and 3 mm wide. Seeds linear-pyram-
idate, brown, mostly 1 mm long.

TYPE.— Panama. Chiriqui: along road between Gualaca and
the Fortuna Dam site at 10. 1 m; NW of Los Planes de Hornito,
elev. 1300 m, 8 Apr. 1980, Antonio 4078 (holotype: CAS!;
isotype: MO).

ADDITIONAL SPECIMENS EXAMINED.— Panama. Chiriqui:
Cerro Hornitos, 40 km NW of Gualaca in cloud forest dom-
inated by Quercus spp., Podocarpus, and Drimys, 26 Jul. 1975,
Mori & Bolten 7486 (CAS). Veraguas: Cerro Tute, ca. 10 km
NW of Santa Fe on ridgetop in cloud forest, 19 May 1975,
Mori 6255 (CAS), 3 Aug. 1975, Mori et al. 7581 (CAS).

DISTRIBUTION.— Known only from cloud for-
ests on Cerro Tute in Veraguas province and
Cerro Hornitos and vicinity in Chiriqui province
at elevations from 1000-2238 m. Flowering
specimens have been collected in April, May,
and July. This species is reportedly (fide Mori
6255) one of the most common trees in an area
about 10 km northwest of Santa Fe on Cerro
Tute.

Graffenrieda bella is most closely related to G.
micrantha (Gleason) L. Wms. which differs in

its longer leaf blades (13-20 cm) with five prom-
inently elevated abaxial nerves, longer petioles
(2.5-5 mm), shorter (2-3 mm) narrowly lanceo-
late petals, and smaller fruiting hypanthia (2-2.5
mm). My circumscription of G. bella includes
two morphological entities representing geo-
graphically distinct populations. The collections
from Chiriqui in western Panama have 5-merous
flowers and foliar margins with bilaterally de-
veloped reflexed auricles about 2 mm above the
petiole/laminar junction. Collections from Vera-
guas in central Panama have 5(-6)-merous flow-
ers and lack the conspicuous foliar auricles. Ex-
cept for these differences, the few available
specimens from each area are identical and ex-
hibit parallel ranges in foliar size, hypanthial and
staminal length, and petal width. More collec-
tions, especially from intervening areas, are
needed to determine whether this bimodal vari-
ation shows a consistent geographical correla-
tion.

Miconia Ruiz & Pavon

As presently circumscribed, Miconia with ap-
proximately 1000 species is the largest genus in
the family and one of the largest among flowering
plants. It is generally characterized by its ter-
minal inflorescences and apically rounded, ob-
tuse, or retuse petals. Miconia occupies a central
position in the complex of genera comprising the
taxonomically difficult tribe Miconieae. Through
its broad neotropical range Miconia displays great
diversity in habit, foliar characters, inflorescence
organization, trichome and androecial mor-
phology, and seed structure. Success in identi-
fying species, however, inevitably requires ex-
amination of anther structure because the current
sectional classification relies heavily on this char-
acter. Miconia is greatly in need of monographic
study to evaluate character variation, assess the
status of satellite groups, and redefine sectional
limits based on character correlations. Some 60
species of Miconia occur in Panama. These rep-
resent all twelve currently recognized sections.
In addition to the species proposed below, sev-
eral new taxa await formal description pending
collection of complete material.

Miconia arboricola Almeda, sp. nov.

(Figure 4)

Caulis scandens primum sicut foliorum subtus
venae primariae inflorescentia hypanthiaque
dense vel modicepilis stipitato-stellatis 0.5-1 mm

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17

FIGURE 4. Miconia arboricola Almeda. A, habit, x ca. 'A; B, representative leaf (lower surface) with enlargement (left), x
ca. '/2; C, stipitate-stellate trichome, x ca. 17; D, mature berry, x ca. 5; E, representative flower showing young hypanthium,
two petals, style, bracteoles, and pedicellar scars, x ca. 7; F, stamens, lateral view (left) and dorsal view (right), x ca. 9; G,
petal, x ca. 7; H, seeds, x ca. 20. (A-C from the holotype; D, F, G from Knapp 5735; E & H from Croat 49151.)

latis indutus. Petioli 8-23 mm longi; lamina 5.5-
11.4 x 3.5-7.1 cm late ovata vel cordata, apice
breviter acuminato basi subcordata vel cordata,
5-7-nervata, fir me coriacea et integra. Panicula
11.5-25 cm longa multiflora; flores 5-meri sub-
sessiles in glomerulis interrupts vel ad ramulo-
rum apices aggregati, bracteolis 1-2.5 x 0.25-

0.5 mm anguste oblongis persist entibus. Hypan-
thium (ad torum) 1.5-2 mm longum; calycis tu-
bus non eminentibus, lobis interioribus 0.5 mm
longis deltoideis, dentibus exterioribus crassis lo-
bos interiores aequantibus. Petala 2-3 x 7.5-2
mm glabra obovata apice retuso. Stamina
isomorphica glabra; fdamenta 2.5 mm longa;

ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA

279

anther arum thecae 1.5 x 0.5 mm oblongo-
cuneatae; connectivum nee prolongatum nee ap-
pendiculatum. Stylus 4 mm glaber; stigma cap-
itatum; semina 0.5 mm galeiformia.

Pendent woody vines or hemiepiphytes ad-
hering to bark of host tree by adventitious roots.
Older cauline internodes terete and ± glabrate.
Distal branchlets, petioles, vegetative buds, ju-
venile leaves, and inflorescences moderately to
copiously covered with ferrugineous stipitate-
stellate trichomes. Leaves of a pair essentially
equal in size; blades coriaceous at maturity, en-
tire, 5.5-11.4 cm long and 3.5-7.1 cm wide,
broadly ovate to cordate, short acuminate api-
cally and subcordate to cordate basally, 5-7-
nerved with a prominulous reticulate network of
secondaries and tertiaries below, mostly glabrate
and vernicose above at maturity or with stipi-
tate-stellate trichomes persisting to varying de-
grees in the furrows created by the impressed
primary nerves, sparsely to moderately stipitate-
stellate below; petioles 8-23 mm long and 2-2.5
mm wide. Inflorescence basically paniculiform
with ultimate branchlets terminating in multi-
flowered congested glomerules; bracts of the
rachis nodes paired, linear-oblong, 2.5-5(-10)
mm long, 1-1.5 mm wide, glabrous adaxially and
stipitate-stellate to glabrate abaxially; bracteoles
3-5 per pedicel, sessile, linear-oblong, acute to
obtuse or rounded apically, 1-2.5 mm long, 0.25-
0.5 mm wide, entire, glabrous adaxially and stip-
itate-stellate abaxially. Pedicels 0.5 mm long but
typically inconspicuous and concealed by con-
gested glomerules. Hypanthia (at anthesis) sub-
cylindric to narrowly campanulate, 1.5-2 mm
long to the torus, moderately to copiously beset
with short stalked-stellate trichomes and occa-
sionally with a sparse scattering of spreading
glandular trichomes on or near basal portions of
the calyx teeth; adaxial rim of the torus minutely
glandular-puberulent. Calyx lobes (on young
fruiting hypanthia), glabrous, hyaline, and erect,
deltoid, 0.5 mm long and 1 mm wide basally;
calyx teeth bluntly subulate, copiously stellate,
adnate to and ± equaling subtending calyx lobes.
Petals 5, glabrous, white, obovate to oblong-ob-
ovate, strongly reflexed, irregularly retuse api-
cally, 2-3 mm long and 1.5-2 mm wide. Stamens
10, isomorphic, filaments straight to somewhat
incurved distally, glabrous, subulate, 2.5 mm
long; anthers 1.5 mm long and 0.5 mm wide
distally, linear-oblong to narrowly cuneate, typ-

ically recurved distally; connective thickened
dorsally but not dilated or prolonged below the
point of filament insertion. Ovary (young fruiting
hypanthia) ca. '/3 to '/4 inferior, 3-celled, coarsely
papillate to farinaceous, caducously glandular-
puberulent along the bluntly fluted stylar collar
that becomes depressed and inconspicuous on
mature fruits. Style straight, glabrous, 4 mm long;
stigma broadly capitate. Berry reportedly blue at
maturity, globose, 3-4 mm long and 3-4 mm in
diameter. Seeds galeiform, white, papillate to
costate-papillate, 0.5 mm long.

TYPE. — Panama. Chiriqui: slope NW of confluence of Rio
Hornito and Rio Chiriqui, ca. 8°44'N, 80°07'W, elev. 1050-
1100 m, 11 Nov. 1980, Stevens 18266 (holotype: CAS!; iso-
type: MO).

ADDITIONAL SPECIMENS EXAMINED.— Costa Rica. Cartago:
Pejibaye, 30 May 1924, Lankester 877 (US). Limon: Cerro
Ur6n, Cordillera Talamanca, 1 Sept. 1898, Pittier 12683 (US).
Panama. Code: hills N of El Valle, E slope and ridges leading
to Cerro Gaital, 8°40'N, 80°07'W, 27 June 1982, Knapp 5735
(MO); Alto Calvario, above sawmill on Continental Divide,
5.2 mi above El Cope, 6 Dec. 1979, Croat 49151 (CAS).

DISTRIBUTION. — This little-collected cloud
forest species ranges from the Caribbean slopes
of the Cordillera Talamanca in south-central
Costa Rica south and east to the hills surround-
ing El Valle de Anton in central Panama at el-
evations from 300-1100 m. Flowering speci-
mens have been collected in June, September,
and November.

Labels on collections of this species describe
it as a pendent woody vine. The notes on Croat
49151 describe it as a hemiepiphyte with the
stem rooted in the ground but closely attached
to a tree trunk. It seems likely that plants of this
species are typically hemiepiphytes that ulti-
mately become scandent vines requiring the sup-
port of host trees. The specific epithet refers to
this discordant habit among species of Miconia.

Miconia arboricola is quite unlike all other
species of Miconia from Central America. In ad-
dition to the unusual habit, it is distinctive in its
coriaceous leaves, stipitate-stellate pubescence,
paniculiform inflorescence, congested, polybrac-
teolate flowers, and sculptured, galeiform seeds.
As pointed out on the label of Pittier 12683, in
Cogniaux's monograph, this species comes clos-
est to M. mollicula Triana and M. cordata Triana
in section Miconia. In foliar shape, internodal
pubescence, and inflorescence structure, M. ar-

280

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17

FIGURE 5. Topobea calophylla Almeda. A, habit, xVr, B, stamens, ventral view, xl; C, hypanthium with floral bracts
removed, x ca. 1 W, D, cauline trichomes, x ca. 4; E, petal, x ca. 1 'A; F, outer floral bract (outer surface), x ca. 1 ; G, inner
floral bract (inner surface), x ca. 1 . (A & D from Nee 9873; B, C & E from Mori & Kallunki 5353; F-G from the holotype.)

ALMEDA: ADDITIONS TO THE MELASTOMATACEAE OF PANAMA

281

boricola resembles these two shrubby species.
They differ most notably from the new species
in having irregularly denticulate leaves, sessile
stellate or stellate-furfuraceous hypanthial pu-
bescence, and a differentiated androecium in
which the larger stamens are inserted opposite
the petals.

Topobea Aublet

The species proposed below represents the first
new Topobea to be described from Panama since
the appearance of Gleason's (1958) summary.
The following description of T. calophylla, the
transfer of T. elliptica to Blakea (also proposed
herein) and the discovery of T. pittieri Cogn. in
Chiriqui province (Cerro Colorado, Folsom &
Collins 1770, CAS) now brings the number of
Panamanian species to nine.

Topobea calophylla Almeda, sp. nov.

(Figure 5)

Frutex epiphyticus. Ramuli primum obscure
quadrangulati demum teretes sicut petioli lami-
narum subtus venae primariae pedicelli bracte-
aeque pilis barbellatis ca. (l-)3-9 mm longis in-
duti. Petioli 2.2-5 cm longi; lamina 14.5-37.5 x
8.6-17.8 cm, elliptico-ovata, apice abrupte cau-
dato-acuminato basi rotundata vel paulo corda-
ta, supra ad maturitatem glabra, subtus modice
setosa (pilis ca. 1-3 mm longis). Flores 6-meri
in foliorum superiorem axillis oppositis plerum-
que 2-4; bracteae liber ae; bracteae exterior es 1.7-
2.3 x 1.5-1.7 cm late ovatae; bracteae interiores
1.6-1.9 x 1.3-1.7 cm elliptico- ovatae. Hypan-
thium (ad torum) 1.1-1.3 cm longum extus dense
strigosum pilis 2-4 mm longis, calycis lobis 14-
17 mm longis triangular ibus. Petala glabra 2 x
1 cm spathulata apice hebeti-acuto vel obtuso.
Filamenta 5 mm longa; anther ae 8 x 1.5 mm
subulatae inter se cohaerentes, poris binis ter-
minalibus; connectivum ad basim dorsaliter mi-
nute calcaratum. Stigma capitellatum vel trun-
catum; stylus 11-14 mm glaber in ovarii collo 5-
6 mm immersus.

Coarse spreading epiphytic shrub. Cauline in-
ternodes ± terete to subquadrangular. Distal
branchlets, vegetative buds, pedicels, and floral
bracts covered with a hirsute indument of rusty
brown barbellate trichomes mostly 3-9 mm long.
Mature leaves firmly chartaceous to coriaceous,
inconspicuously dentate, 14.5-37.5 x 8.6-17.8
cm, elliptic-ovate, abruptly caudate-acuminate

apically, rounded to subcordate basally, 5-7-
nerved with conspicuous elevated secondaries,
glabrous above, moderately hirsute below with
barbellate trichomes mostly 1-3 mm long; pet-
ioles 2.2-5 cm long and 3-5 mm diam. Flowers
6-merous, erect to widely spreading, paired or
borne in clusters of three or four in axils of the
distal branches; pedicels 2.8-4 cm long. Floral
bracts foliaceous, entire, 3-5 -nerved free basally,
each pair closely subtending one another or sep-
arated on the pedicel by a distance of 3-4 mm,
outer bracts 1 .7-2.3 x 1.5-1 .7 cm, broadly ovate,
acuminate apically; inner bracts 1.6-1.9 x 1.3-
1.7 cm, elliptic-ovate, acute to acuminate api-
cally. Hypanthia (at an thesis) campanulate, 1.1-
1.3 cm long to the torus, copiously appressed-
strigose with barbellate trichomes 2-4 mm long.
Calyx lobes lance-triangular, often involute to
uncinate apically when dry, entire, hirsute to hir-
tellous, 14-17 mm long and 5-6 mm wide ba-
sally between sinuses. Petals glabrous, reportedly
pink, ± spathulate but acute to obtuse apically,
entire to sparingly glandular-ciliate, 2 cm long
and 1 cm broad distally. Stamens isomorphic
and strongly declined to one side of the flower,
filaments glabrous, connivent, 5 x 1.5 mm; an-
thers linear-subulate, 8 x 1.5 mm, laterally co-
herent but free distally, strongly incurved at an-
thesis, each anther tipped with two somewhat
dorsally inclined confluent pores; connective
thickened dorsally near the point of filament in-
sertion into a blunt callosity. Style ± straight,
glabrous, 1 1-14 mm long and 1 mm wide; stigma
capitellate to truncate, the surface appearing
somewhat crateriform. Ovary inferior, 6-celled,
distended apically into a glandular puberulent
fluted cone and stylar collar mostly 7-9 mm long.
Mature berry not seen.

TYPE. — Panama. Veraguas: 5 mi W of Santa Fe on road past
Escuela Agricola Alto Piedra on Pacific side of divide, elev.
800-1200 m, 18 Mar. 1973, Croat 23000 (holotype: CAS!;
isotype: MO).

ADDITIONAL SPECIMENS EXAMINED.— Panama. Veraguas: 6-
7 km W of Santa Fe on new road past agriculture school, 1 8
Feb. 1974, Nee 9873 (CAS); NW of Santa Fe, 4.2 km from
Escuela Agricola Alto de Piedra, 25 Feb. 1975, Mori & Kal-
lunki 4833 (CAS); NW of Santa Fe, 2.7 km from Escuela
Agricola Alto de Piedra along stream on road to Calovebora,
30 Mar. 1975, Mori & Kallunki 5353 (CAS).

DISTRIBUTION.— Known only from montane
forests NW of Santa Fe in Veraguas province at
elevations of 800-1200 m. Available material,

282

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 17

all of which is in flower, was collected in February
and March.

Like so many epiphytic species in the Blakeae
that grow in wet forests, this species has a re-
stricted distribution and does not appear to be
closely related to any described taxon in Topo-
bea. As emphasized by the specific epithet, T.
calophylla has large, handsome leaves that are
glabrous above and moderately hirsute below. It
is also striking because of the copious indument
of brown barbellate trichomes that gives distal
branches, floral bracts, and hypanthia a coarse
bristly appearance.

Among the species of Topobea recorded for
Panama by Gleason (1958), this species also dif-
fers in the following combination of characters:
its leaves are inconspicuously dentate; the inner
and outer floral bracts are separate to the base,
copiously pubescent abaxially, and essentially
equal in length; and the ovary is elaborated api-
cally into a prominent glandular-puberulent,
fluted cone crowned by several setiform lobules
that envelop the style basally for 5-6 mm.

No information is available on the size of in-
dividual plants in this species. Judging from fo-
liar size and the coarseness of its branches, I
suspect that T. calophylla can become an epi-
phyte of massive proportions comparable to To-
pobea durandiana Cogn. and Blakea tuberculata
Donn. -Smith, both of which can obscure and

overtake the crowns of their host trees. Casual
collectors unfamiliar with the propensity for epi-
phytism in Topobea and Blakea frequently de-
scribe the epiphytic species in these genera as
trees. Labels for all except one of the few avail-
able collections of T. calophylla indicate that it
is an epiphyte. Although Nee 9873 is described
as a small tree, additional field study is needed
to confirm this observation.

ACKNOWLEDGMENTS

I am grateful to Terry Bell for preparing the
line drawings, to Colleen Sudekum for technical
assistance, and to J. J. Wurdack for critical re-
view of the manuscript. I also thank curators of
the following herbaria who provided special loans
and/or made their facilities available for study:
BM, CR, DUKE, F, MO, US.

LITERATURE CITED

ALMEDA, F. 1974. A new epiphytic Blakea (Melastomata-

ceae) from Panama. Brittonia 26:393-97.
. 1 980. Central American novelties in the genus Blak-
ea (Melastomataceae). Rhodora 82:609-15.
COGNIAUX, C. A. 1891. Melastomaceae. Pages 1-1256. In

A. and C. de Candolle, eds., Monographiae phanerogama^

rum, vol. 7. G. Masson, Paris.
GLEASON, H. A. 1958. Melastomataceae. In R. E. Woodson,

Jr. and R. W. Schery, Flora of Panama. Ann. Missouri Bot.

Gard. 45:203-304.
WURDACK, J. J. 1973. Melastomataceae. In T. Lasser, ed.,

Flora de Venezuela 8:1-819.

CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, California 94 1 1 8

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 18, pp. 283-300, 10 figs., 1 table. December 11, 1984

MARINE AND FRESHWATER STINGRAYS (DASYATIDAE) OF

WEST AFRICA, WITH DESCRIPTION

OF A NEW SPECIES

By
Leonard J. V. Compagno

Tiburon Center for Environmental Studies, San Francisco State University,
P.O. Box 855, Tiburon, California 94920

and
Tyson R. Roberts

California Academy of Sciences, Golden Gate Park, San Francisco, California 94118

ABSTRACT: Three genera and 12 species of stingrays, all members of the family Dasyatidae, are known from
West Africa. A key is provided for their identification. The two most common species in shallow coastal
waters (including lagoons, estuaries, and river mouths) are the closely related Dasyatis margarita (Giinther,
1870) and Dasyatis margaritella new species, which differ in disc shape, meristic features, and adult size.
Two species occur mainly or exclusively in fresh water: Dasyatis garouaensis (Stanch and Blanc, 1962) and
Dasyatis ukpam (Smith, 1863). Dasyatis garouaensis, a small, thin-bodied species previously known only from
the lower Niger and a tributary, the Benue, is reported from Lagos and the Cross River. It is closely related
to D. margarita and />. margaritella. Dasyatis ukpam, a large thick-bodied species with a vestigial sting
previously known only from fetal specimens obtained at Old Calabar (without precise information on habitat),
is now reported from the Ogooue and the lower Zaire or Congo rivers. It is perhaps related to the genus
Urogymnus.

INTRODUCTION Ogooue basin in 1978. We have also identified

This study was undertaken to clarify the sys- a specimen of this species collected in the lower

tematics, distribution, and relationships of West Zaire (Congo) River in 1937.

African freshwater stingrays. Although widely The other West African freshwater dasyatid,

distributed and familiar to local fishermen, sting- D. garouaensis, was described originally as a

rays inhabiting the larger rivers of West Africa species of Potamotrygon, a genus of the otherwise

are poorly known scientifically. There are at least exclusively Neotropical freshwater family Po-

two species. One, Dasyatis ukpam, was de- tamotrygonidae. Evidence that it is actually a

scribed more than a century ago, but the two member of the family Dasyatidae was advanced

fetal type-specimens obtained at Old Calabar by Thorson and Watson (1975). Our own ob-

lacked precise habitat data, and the species was servations fully support this conclusion. Reid and

not reported again or recognized as valid until Sydenham (1979) suggested that D. garouaensis

the junior author obtained specimens in the may be identical with the coastal species D. mar-

[283]

284

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18

garita. Our studies, however, indicate that it is
a valid species, albeit a close relative of D. mar-
garita and the previously undescribed D. mar-
garitella. Dasyatis garouaensis was known only
from the Benue and lower Niger, but the junior
author collected a specimen in the Cross River,
Cameroun, in 1980, and we have also identified
a specimen from Lagos, Nigeria (habitat un-
known). So far as we have been able to deter-
mine, D. margarita and D. margaritella occur
only in marine and estuarine habitats.

The key below should permit identification of
all stingrays now known from West Africa. Fol-
lowing the key we present a definition of the
genus Dasyatis and detailed descriptions of D.
margarita, D. margaritella, D. garouaensis, and
D. ukpam.

MATERIALS AND METHODS

This paper is based on material deposited in
the American Museum of Natural History
(AMNH); British Museum (Natural History)
(BMNH); California Academy of Sciences (CAS
and CAS-SU); Institut fur Seefischerei, Hamburg
(ISH); Museum National d'Histoire Naturelle,
Paris (MNHN); Musee Royale de 1'Afrique Cen-
trale, Tervuren, Belgium (MRAC); and Smith-
sonian Institution (USNM).

Disc width (measured at widest point) is our
standard measure of size, and proportional mea-
surements (unless otherwise indicated) are ex-
pressed as percent of disc width. Definitions or
explanations of some other terms are as follows:

Disc length — midline measurement from
snout-tip to a transverse line parallel to poste-
riormost extension of pectoral fins

Disc depth— greatest height or depth of disc
(usually at scapulocoracoid)

Preorbital length— midline measurement from
snout-tip to a transverse line parallel to anterior
margin of eyes

Prenarial length— midline measurement from
snout-tip to a transverse line parallel to anterior
border of nostrils

Prebranchial length— midline measurement
from snout-tip to a transverse line parallel to
opening of first gill slits

Head length — midline measurement from
snout-tip to a transverse line parallel to opening
of fifth gill slits

Pectoral fin inner margin— from posterior in-
sertion to posteriormost extension of pectoral fin

Pelvic fin span— distance between apices of
pelvic fins when maximally extended

Upper and lower tooth rows — maximum
number of tooth rows across upper and lower
jaws

Vertebral counts in stingrays are complicated
by the extraordinary specialization of the ver-
tebral column as a support for the pectoral fins,
and by its termination in an elongate tail, which
is frequently damaged. Anteriorly the column is
fused into two synarcuals incorporating a vari-
able number of centra. In Dasyatidae the ante-
riormost 23-40 vertebrae are incorporated into
the first synarcual. In most of these vertebrae the
centra are completely fused, but their number
can be determined by counting the spinal nerve
foramina. The second synarcual frequently is
separated from the first by a small number of
intersynarcual vertebrae; in most of the Dasyatis
herein reported, however, there is only a joint
between the two synarcuals. In the second syn-
arcual the centra, although fused, retain their form
and are readily counted in radiographs. Some-
times the posteriormost centrum in the second
synarcual is sharply distinguished from the
monospondylous trunk centra succeeding it. In
specimens in which the end of the second syn-
arcual cannot be determined, we give a combined
count of second synarcual plus monospondylous
trunk vertebrae. This is usual in late fetal or new-
born specimens with poor calcification and in
heavily denticulated specimens in which this
portion of the vertebral column is obscured in
radiographs (e.g., in D. ukpam). Posteriorly the
vertebral column ends in a long series of diplos-
pondylous tail centra followed by a highly flex-
ible, unsegmented rod (apparently consisting of
the notochord and a heavily calcified notochor-
dal sheath). The monospondylous and diplos-
pondylous sections of the vertebral column are
usually sharply demarcated in radiographs.

For terminology and illustrations of dasyatid
clasper morphology see Compagno and Roberts
(1982).

Family DASYATIDAE Jordan, 1888

We follow Bigelow and Schroeder (1953) in
restricting Dasyatidae to the whiptailed sting-
rays, and tentatively recognize the following gen-
era: Dasyatis, Himantura, Hypolophus, Taeni-
ura, Urogymnus, and Urolophoides (see also
Compagno and Roberts 1982).

COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS

285

FIGURE 1. Disc shape in West African Dasyatidae. (a) Dasyatis violacea (trapezoidal); (b) Dasyatis centroura (diamond-
shaped); (c) Taeniura grabata (circular); (d) Urogymnus asperrimus (oval).

KEY TO DASYATIDAE OF WEST AFRICA

la. Disc oval (Fig. Id); tail without dermal

folds; sting invariably absent

Urogymnus

africanus (Bloch and Schneider, 1801)

Ib. Disc variable in shape, tail with dermal
fold or folds, sting usually present (ab-
sent in some Dasyatis ukparri) 2

2a. Ventral tail fold extending to end of tail;

disc circular (Fig. Ic) Taeniura

grabata (E. Geoffrey Saint-Hilaire, 1817)

2b. Ventral tail fold ending far anterior to
end of tail (Dasyatis) 3

3a. Disc trapezoidal or diamond-shaped (Fig.

la-b) 4

3b. Disc oval or circular 9

4a. Disc trapezoidal, anterior margin broad-
ly rounded, snout not projecting as an
angular lobe from disc (Fig. la); upper

and lower surfaces of disc dark

D. violacea (Bonaparte, 1832)

4b. Disc diamond-shaped, anterior margin
angular, snout projecting as an angular
lobe from disc (Fig. Ib); lower surface of
disc light 5

5a. Upper surface of disc with a dark retic-
ular pattern; ventral tailfold very short,

about twice length of sting

D. marmorata (Steindachner, 1892)

5b. Upper surface of disc plain; ventral tail-
fold long, much more than twice sting
length 6

6a. Entire dorsal surface of disc covered with
small denticles; no middorsal row of en-
larged denticles or thorns; adults with
over 100 rows of teeth in each jaw; disc
very broad, about 1.5 times as wide as
long in adults D. rudis (Gunther, 1870)

6b. Dorsal surface of disc only partially cov-
ered with small denticles, along middle
of back, or naked except for a middorsal
row of enlarged denticles or thorns; adults
with much less than 100 rows of teeth in
each jaw; disc narrower, 1.0-1.3 times
as wide as long 7

7a. Anterior margin of disc anterior to spi-
racles nearly straight behind snout-tip,
with tip projecting; posterior parts of pel-
vic fins projecting well rearward beyond

rear tip of pectoral fins

D. pastinaca (Linnaeus, 1758)

7b. Anterior margin of disc anterior to spi-
racles slightly concave behind snout-tip,
with tip not conspicuously projecting;
posterior parts of pelvic fins extending
slightly behind rear tips of pectoral fins
8

8a. Ventral tailfold high, about as deep as
tail above it; a dorsal ridge present on
tail behind sting; disc and tail in large
juveniles and adults without enlarged,
heavy, broad-based denticles, but with
moderately enlarged middorsal and

scapular denticles only D. ameri-

cana (Hildebrand and Schroeder, 1928)

286

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18

8b. Ventral tailfold low, half as deep as tail
above it; no dorsal ridge on tail behind
sting; disc and tail in large juveniles and
adults with scattered enlarged, heavy,
broad-based denticles, forming a mid-
dorsal row at center of disc and tail

D. centroura (Mitchill, 1815)

9a. Anterior margins of disc broadly convex
anterior to spiracles, with tip of snout
not projecting from them; dorsal disc
surface in juveniles to subadults com-
pletely covered with denticles, with flat-
tened large denticles on midbelt, small
pointed denticles laterally, and large,
conical, erect, sharp-cusped denticles on
thorns scattered on disc and tail base;
sting reduced or absent; no dorsal keel
on tail; base of tail circular in cross sec-
tion; ventral surface of disc light with a

broad dusky marginal band

D. ukpam (Smith, 1863)

9b. Anterior margins of disc concave ante-
rior to spiracles, with tip of snout con-
spicuously projecting from them; dorsal
disc surface partially naked, with a mid-
belt of flattened denticles and often a
midscapular pearl spine, or naked; no
large conical thorns on disc and tail base;
sting large; a low dorsal keel on tail be-
hind sting; base of tail horizontally oval
in cross section; ventral surface of disc
light without a broad dusky marginal
band 10

1 Oa. Back flattened, without an enlarged mid-
scapular pearl spine (sometimes a row of
up to 4 moderately enlarged flattened
spines in its place); midbelt of flattened
denticles reduced or absent, disc some-
times entirely naked; snout long, preor-
bital length 2.8-3.2 times interorbital
space (down to 2.3 in late fetuses or new-
born specimens) and 27-32% of disc
width; disc very flat, thickness at scap-
ulocoracoid 8-11% (usually less than
1 1%) of disc width; lateral prepelvic pro-
cesses of pelvic girdle greatly expanded

D. garouaensis

(Stauch and Blanc, 1962)

1 Ob. Back somewhat arched, with an enlarged
midscapular pearl spine; midbelt of flat-
tened denticles well developed in large

juveniles and adults; snout shorter,
preorbital length 1.5-2.4 times interor-
bital space and 1 9-26% of disc width;
disc thicker, 1 1-15% of disc width over
scapulocoracoid; lateral prepelvic pro-
cesses slightly expanded 11

1 1 a. Upper jaw strongly undulated, with teeth
greatly enlarged on prominent lateral
projections; teeth less numerous, in 26-
29/31-34 rows; snout more broadly
pointed; pearl spine usually larger and
circular, length about 5-6 mm; pectoral
radials 133-135; size larger, adults to 65
cm D. margarita (Gunther, 1870)

lib. Upper jaw moderately undulated, with
teeth moderately enlarged on low lateral
projections; teeth more numerous, in 36-
42/38-50 rows; snout usually more
acutely pointed; pearl spine usually
smaller and often axially oval, length 2-
4 mm; pectoral radials 116-127; size

smaller, adults to 26 cm

D. margaritella new species

Dasyatis Rafinesque, 1810

Dasyatis RAFINESQUE, 1810:16 (type-species Dasyatis ujo Ra-
finesque, 1810 [=Rajapastinaca Linnaeus, 1 758], by mono-
typy).

For full generic synonymy of Dasyatis see Bigelow and
Schroeder(1953).

DIAGNOSIS. — Dasyatidae with disc circular,
oval, trapezoidal, or diamond-shaped (Fig. 1), its
dorsal surface smooth or variably covered with
small, flat or prickle-like denticles; large, sharp,
spine- or plate-like denticles present or absent
on dorsal surface; snout angular or broadly
rounded, its projecting tip variably developed;
pectoral fins rounded or angular; pelvic bar mod-
erately arched; tail long, slender, with dorsal and
ventral folds or ventral folds only; ventral fold
not reaching tip of tail; sting usually present (re-
duced or absent in Dasyatis ukpam). Teeth small,
rhomboidal, thin-crowned.

Dasyatis as here recognized is a large, heter-
ogeneous assemblage of about 33 species and
may be polyphyletic. Dasyatis margarita, D.
margaritella, and D. garouaensis are not far re-
moved morphologically from the generic type-
species D. pastinaca. Dasyatis ukpam, however,
is distinct, approaching Urogymnus Muller and
Henle, 1837 in general morphology, heavy den-

COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS

287

FIGURE 2. (a) Dasyatis margarita, lectotype, 200-mm immature female, West Africa (BMNH 1865.7.4:1); (b) Dasyatis
margaritella, 226-mm mature male, Conakry, Guinea (ISH 183/63).

ticulation, and sting reduction. Urogymnus
species invariably lack the sting, while specimens
of D. ukpam either lack the sting or have a very
small one. Smith (1863) noted that D. ukpam
seemed intermediate between Urogymnus and
Trygon (=Dasyatis), at least in the nature of its
sting, but included it in Hemitrygon Miiller and
Henle, 1837 (=Dasyatis) because it has a short
ventral tail fold and no dorsal tailfold. We retain
D. ukpam in Dasyatis pending modification of
the limits of Dasyatis and other dasyatid genera.
The species is readily distinguished from known
Urogymnus species in having a ventral tailfold,
much longer tail (less than 1.5 times disc width
in Urogymnus), a less thick, more circular disc,
darker dorsal coloration (dorsal surface pale
brown in all Urogymnus examined), a dark mar-
ginal band on ventral surface of disc, smaller flat
denticles on dorsal surface of disc, and in some
specimens a small sting.

Dasyatis margarita (Giinther, 1870)
(Figure 2a)

Trygon margarita GUNTHER, 1870:479 (type-locality West Af-
rica).
Dasyatis sp. BLACHE ET AL., 1970:53, fig. 117.

MATERIAL EXAMINED. — BMNH 1865.7.4:1, 200-mm im-
mature female, West Africa (formerly syntype ofD. margarita;
designated lectotype below); USNM 222589, 1 30-mm late fetal
or newborn male, Sierra Leone; BMNH 1930.3.24:3, 212-mm
immature male, Accra, Ghana; BMNH 1936.8.20:2-3, 216-
mm and 315-mm immature females, Lagos, Nigeria; BMNH
1899.2.20:35, 206-mm immature female, Banana, Congo Riv-
er mouth, Zaire; AMNH 40408, 235-mm female, Angola.

LECTOTYPE DESIGNATION.— In the original description of D.
margarita Glinther ( 1 870:479) listed two specimens from West
Africa without indicating either as holotype:

a. Disk 8'/2 inches long, tail 19 inches. Purchased of Mr. J.
Wood.

b. Young. From the collection of the Zoological Society.

These two specimens are therefore syntypes, but our studies
indicate they are not conspecific. Specimen a is BMNH 1 865.7.4:
1 , a 200-mm immature female with a single, large, round pearl

288

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18

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spine; specimen b apparently is BMNH 1852.8.30:23, a 1 16-
mm fetal or newborn male with two relatively small oval pearl
spines. Gttnther's description clearly is based mainly on spec-
imen a, which we therefore designate as lectotype. Specimen
b belongs to D. margaritella.

DIAGNOSIS.— A large marine and estuarine
Dasyatis, 65 cm or more, most closely related to
the much smaller marine and estuarine species
D. margaritella and to the extremely flat fresh-
water species D. garouaensis. Pearl spines always
present and very large (much smaller in D. mar-
garitella and absent or represented by 2-3 low
denticles in D. garouaensis). Denticles on dorsal
surface of disc smooth, restricted to middle por-
tion.

Disc oval, moderately flat, its medial lobe
broad-based and moderately exserted, its an-
terolateral margin somewhat concave; disc depth
13.0-14.5% of disc width, disc width 0.9-1.1
times disc length; snout moderately long, preoral
length 22-24% of disc width; dorsal surface of
disc naked in young (except for pearl spine), but
in older individuals (larger than about 200 mm)
middle third of disc covered with small, circular,
flat denticles and tail covered with small prickles;
no enlarged thorns on disc or tail; a massive,
usually circular, button-shaped pearl spine on
back, 4.8-5.5 mm long; usually a single large
slender sting on tail, its length 22% of disc width
in newborn; eyes moderately large and somewhat
elevated, eyeball length 1.4-2.3 times in inter-
orbital space, interorbital space 1 .5-2.0 in preor-
bital length; spiracles moderately large and flat-
tened; floor of mouth with 5 elongate papillae;
total tooth rows 26-30/31-34; pelvic fins short,
anterior margins 1 8-2 1 % of disc width; tail evenly
tapering to a slender whiplash behind sting, but
broad opposite and in front of it, its length when
intact about 1 .8-2.5 times disc width; base of tail
horizontally oval and depressed in section; ven-
tral tailfold moderately high, dorsal tailfold re-
duced to a low keel behind sting; disc and pelvic
fins gray-brown above, without spots or prom-
inent markings, uniformly pale or whitish below;
intestinal valve turns 12-14; total pectoral ra-
dials 133-135; total vertebral centra 130, and
total vertebral segments 162; vertebral centra ex-
tending behind origin of sting; a moderately large
stingray, exceeding 65 cm.

Proportional measurements (as percent of disc
width) and counts are given in Table 1 . Preoral
length 2.5-3.0 times internarial width and about
1 .0 times width between first gill slits. Snout fair-

290

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18

ly broad, angle in front of eyes 123°. Spiracle
length 1.0-1.3 times eyeball length, 0.8-0.9 times
internarial width, and 1.8-2.0 times in distance
between fifth gill slits. Nasal curtain with a fringed,
weakly trilobate posterior margin. Mouth weakly
arched, midline of lower jaw with a prominent
indentation; a shallow to deep, curved groove
extending posteriorly from posterior nasal flap
around corners of mouth. Floor of mouth with
5 papillae, including a transverse row of one small
medial and two large elongate paramedial pa-
pillae, and two smaller lateral papillae near ends
of dental bands.

Teeth in quincunx pavement, with crowns
closely adjacent to one another. Teeth similar in
upper and lower jaws, upper and lower jaw teeth
about equally large at symphysis, varying con-
tinuously lateral to synphysis, largest and most
elongate at or near symphysis and smallest and
least elongate at mouth corners. Upper dental
band with a prominent knob of considerably en-
larged teeth at symphysis, separated from simi-
lar, very large knobs at either side by depressions
with smaller teeth; lower dental band with a well-
marked symphyseal depression into which sym-
physeal knob of upper jaw fits, and a pair of
prominent lateral knobs fitting into depressions
in upper dental band.

Dorsal surface naked except for pearl spine in
young below about 200 mm, but in larger spec-
imens moderately large, heart-shaped or circular
denticles form a middorsal belt of denticles on
disc; lateral parts of disc naked. Larger individ-
uals with small conical prickles on dorsal surface
of tail behind sting.

Neurocranium, observed in radiographs, sim-
ilar to that of Himantura signifer as described
by Compagno and Roberts (1 982) but with long-
er, more ovate nasal capsules.

Pelvic girdle (Fig. lOe) broadly arched, semi-
circular, and relatively narrow, with a medial
angle but without a medial prepubic process. Lat-
eral prepubic processes low, rounded, lobate, and
not greatly expanded; iliac processes well devel-
oped; ischial processes short. Four obturator fo-
ramina present.

Claspers not studied in detail, but morpholog-
ically similar to those of D. garouaensis.

Dasyatis margaritella, new species

(Figures 2b, 3)

Trygon margarita GUNTHER, 1870:479 (in part).
Dasyatis margarita BLACHE ET AL., 1970:53, fig. 1 16.

HOLOTYPE.-CAS-SU 68915, 190-mm male, Mbode, Cam-
eroun, A. I. Good, 7 December 1 940.

PARATYPES.— CAS-SU 68916, 169-mm male and 180-mm
female, same collection data as holotype.

ADDITIONAL MATERIAL EXAMINED BUT NOT INCLUDED IN
TYPE-SERIES. -BMNH 1852.8.30.23, 116-mm immature male,
West Africa, apparent syntype of Trygon margarita Gttnther,
1870; USNM 222590, 4 immature males, 109-139 mm, 2
females, 1 32-1 64 mm, and USNM 222593, 3 immature males,
129-141 mm, 146 mm female, both lots from Guinea-Bissau;
ISH 141-142/62, 3 immature males, 135-196 mm, 201-mm
adolescent male, 228-mm adult male, 3 females, 149-222 mm,
Guinea; ISH 1 83/63, 226-mm adult male and 248-mm female,
Conakry, Guinea; ISH 295/59, 167-mm immature male, Du-
breka, Guinea; USNM 222591, 145-mm female, Liberia;
BMNH 1920.8.12.1, 262-mm female, Sierra Leone; USNM
222591, 149-mm female, Liberia; BMNH 1914.1 1.2.75, 133-
mm female, Lagos, Nigeria; BMNH 1937.4.19.4, 208-mm adult
male, Lagos Lagoon, Nigeria; AMNH 41515, 204-mm female,
Congo River mouth.

DERIVATION OF N AME. — margaritella, dimin-
utive of Latin margarita, f. "pearl"; for the smaller
size of this stingray and of its pearl spine, as
compared to D. margarita.

DIAGNOSIS.— D. margaritella is a marine
species and the smallest dasyatid in West Africa.
Males sexually mature at 208 mm or less; largest
known specimen a 262-mm female. Most similar
to the marine species D. margarita, which attains
over 600 mm, has a larger pearl spine, and more
numerous pectoral radials( 133- 135 vs. 1 16-127).
It is also similar to D. garouaensis, a freshwater
species with a much flatter disc and a longer
snout.

Disc oval, moderately flat. Medial lobe broad-
based and exserted, its anterior margin concave.
Disc depth 1 1 .0-1 5.5% of disc width, disc width
0.9-1 . 1 times disc length; snout moderately long,
preoral length 25-27% of disc width; dorsal sur-
face of disc naked in young (except for pearl spine),
but above about 130-140 mm middle third of
disc covered with small heart-shaped or circular
flat denticles and tail posterior to sting with small
prickles; no enlarged thorns on disc or tail; a
moderately large, usually longitudinally oval pearl
spine on back, 2.4-4. 1 mm long; usually a single
large slender sting on tail, 24-32% of disc width;
eyes moderately large and somewhat elevated,
eyeball length 1.2-1.9 times in interorbital space,
interorbital space 1.7-2.4 in preorbital length;
spiracles moderately large and flattened; floor of
mouth with 5 elongated papillae; total tooth rows
35-43/38-50; pelvic fins short, anterior margins
1 3-22% of disc width; tail evenly tapering to a
slender whiplash behind sting, but broad oppo-

COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS

291

FIGURE 3. Dasyatis margaritella, holotype, 190-mm immature male, Mbode, Cameroun (CAS-SU 68915).

site and in front of it, its length from vent to tip
when intact about 2.5-3.4 times disc width; base
of tail horizontally oval and depressed in section;
a moderately high ventral tailfold present but
with only a low keel on the dorsal surface of the
tail behind the sting; disc and pelvic fins gray-
brown above, without spots or prominent mark-
ings, white below and without a marginal dark
band; intestinal valve turns 13-14; total pectoral
radials 1 16-127; total vertebral centra 1 16-128,
and total vertebral segments 139-151; vertebral
centra extending behind origin of sting. A small
stingray, probably not exceeding 30 cm.

Proportional measurements (as percent of disc
width) and counts are given in Table 1 . Preoral
length 2.5-3.3 times internarial width and 1.1-
1 .3 times width between first gill openings. Snout
fairly narrow, angle in front of eyes 113-119°.
Spiracle length 0.7-1 . 1 times eyeball length, 0.6-
0.9 times internarial width, and 1.8-2.5 times in
distance between fifth gill openings. Nasal cur-
tain with a fringed, nearly straight or weakly tri-
lobate posterior margin. Mouth weakly arched,
midline of lower jaw with a prominent inden-
tation; a shallow-to-deep, curved groove extend-
ing posteriorly from posterior nasal flap around

corners of mouth. Skin on ventral surface of low-
er jaw more or less corrugated and papillate. Pal-
ate behind fringed maxillary valve with three
strong, short ridges, a medial longitudinal ridge
and a pair of diagonal lateral ridges. Floor of
mouth with 5 oral papillae, including a trans-
verse row of one smaller medial and two larger
elongated paramedial papillae at midline of
mouth, and two smaller lateral papillae near ends
of dental bands.

Upper jaw with about 6, lower jaw with about
8 functional series of teeth. Teeth in quincunx
pavement, close-set, with crowns closely adja-
cent to one another. Teeth similar in upper and
lower jaws; uppers about as large as lowers at
symphysis, varying continuously lateral to sym-
physis; teeth largest and longest relative to width
at or near symphysis and smallest and shortest
at mouth corners. Upper dental band with a
prominent knob of slightly enlarged teeth at sym-
physis, separated from similar knobs at either
side by depressions with smaller teeth; lower
dental band with a well-marked symphyseal
depression into which symphyseal knob of upper
jaw fits, and a pair of prominent lateral knobs
that fit into depressions in upper dental band.

292

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18

FIGURE 4. Dasyatis garouaensis, 340-mm mature male, Benue River at Makurdi, Nigeria.

Teeth of both jaws with low, broad-tipped, cusp-
less, rhomboidal crowns that have a transverse
blunt keel or carina, reticulating ridges on their
labial sides, a strong basal ledge, and concave,
inset basal groove separating crown and root.
Roots moderately large, fairly low, pedicellate,
bilobed basally, with a transverse groove and
nutrient foramen. Teeth of adult males some-
what longer than in females, but without elon-
gated cusps.

Dorsal surface naked in young below about
1 30-1 40 mm, except for pearl spine, but in larger
specimens moderately large, heart-shaped or cir-
cular denticles form a middorsal belt of denticles
on disc; lateral parts of disc naked. Also, small
conical prickles on dorsal surface of tail behind
sting in larger individuals.

Neurocranium, observed on radiographs, sim-
ilar to that ofHimantura signifer as described in
Compagno and Roberts (1982), but with longer,
more ovate nasal capsules.

Pelvic girdle (Fig. lOf) broadly arched, semi-
circular, relatively narrow, with a medial angle
but without a medial prepubic process. Lateral
prepubic processes low, rounded, lobate, not

greatly expanded; iliac processes well developed;
ischial processes short. Four obturator foramina
present.

Claspers not studied in detail, but similar mor-
phologically to those of Dasyatis garouaensis.

Dasyatis garouaensis (Stauch and Blanc, 1962)

(Figures 4-5)

Potamotrygon garouaensis STAUCH AND BLANC, 1962:166, fig.
1-4 (type-locality Benue River at Malape, Cameroun); DA-
GET AND STAUCH, 1963:85-107 (reference); CASTEX, 1967:
167-176 (discussion).

Dasyatis sp. KREFFT, 1968:70, pi. 6 ("Rafin Kunama, Neben-
fluss des Nigers etwa 300 Meilen oberhalb der Mlindung").

Dasyatis garouaensis CASTELLO, 1973:67 (placed in Dasyatis);
THORSON AND WATSON, 1975:701-712 (placed in Dasyatis,
size, range, physiology, additional specimens reported from
Niger and Benue rivers of Nigeria and Cameroun); REID AND
SYDENHAM, 1979:46, 54-55 (possibly synonymy with D.
margarita, range in Benue River system); COMPAGNO AND
ROBERTS, 1982:321 (reference).

MATERIAL EXAMINED. -MNHN 1962-303, holotype, 202-
mm female; MNHN 1962-304, 68-mm male and 78-mm fe-
male fetuses; MNHN 1 967-44 1 , 263-mm female; MNHN 1 967-
440, 300-mm adult male; MNHN 1967-439, 340-mm adult
male; all from upper Benue River, Cameroun; CAS 49147,
342-mm adult female, Cross River, 5-1 0 km downstream from
Mamfe, Cameroun; CAS 53108, 311-mm female, 340-mm

COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS

293

adult male; Thorson collection, uncat, 238-mm female (ra-
diograph only); Benue River at Makurdi, Nigeria; BMNH
1949.10.24.1, 255-mm adult male, Lagos, Nigeria.

DIAGNOSIS.— A moderately large and thin-
bodied freshwater dasyatid. Largest known spec-
imen a 342-mm adult female. Disc flatter than
in any other West African dasyatid, its depth
only 8.7-1 1 .0% of disc width (vs. 1 1 .0% or more
in other species). Denticles on dorsal surface of
disc highly variable, sometimes absent, always
restricted to central portion. Pearl spine absent
or represented by 2-3 low denticles. Most similar
to the small marine species D. margaritella and
the larger marine species D. margarita, both of
which have well-developed pearl spines and
shorter snouts.

Disc oval, very flat. Medial lobe fairly broad-
based and elongated, its anterior margin straight
or concave; disc width 0.9-1 .0 times disc length;
snout moderately long, preoral length 29-32% of
disc width; dorsal surface of disc either entirely
naked or with scattered, small, heart-shaped or
circular flat denticles on its middle, no pearl spines
(a few slightly enlarged, flattened, heart-shaped
denticles occasionally present in midscapular
area), and no enlarged conical denticles on disc
and tail; a single, large, slender sting present on
tail; sting 22.6-26.3% of disc width in adults;
eyes small and hardly elevated, eyeball lengths
1.3-2.0 times in interorbital space; interorbital
space, 2.4-3.2 times in preorbital length; spira-
cles moderately large and flattened; floor of mouth
with 5 elongated papillae; total tooth rows 32-
40/37-45; pelvic fins short, their anterior mar-
gins 20-25% of disc width; tail rapidly tapering
to a slender whiplash behind sting but broad op-
posite and in front of it, its length from vent 2.5-
3.5 times disc width when intact; base of tail
horizontally oval and depressed in section; a
moderately high ventral tailfold present but with
only a low keel on dorsal surface of tail behind
sting; disc and pelvic fins medium gray or gray-
brown above, without spots or prominent mark-
ings, white below and without dark margins; tail
darker or blackish, mottled, lighter below, un-
derside of base white; intestinal valve turns 10;
total pectoral radials 122-125; total vertebral
centra 120-131 and total segments about 149-
1 62; vertebral centra extending posterior to sting
origin.

Proportional measurements (as percent of disc
width) and counts are given in Table 1 . Preoral

length 4.0-6.0 times internarial width and 1.2-
1.9 times width between first gill slits. Snout
moderately broad, angle in front of eyes 110-
1 16°. Spiracle length 0.9-1.2 times eyeball length,
0.8-1 .2 times internarial width and 1 .8-2. 1 times
in distance between fifth gill slits. Internal gill
openings with close-set transverse ridges on gill
arches, apparently serving as gill rakers. Nasal
curtain with a fringed, concave posterior margin.
Mouth nearly straight, midline of lower jaw
slightly concave. A groove extending posteriorly
from posterior nasal flap around mouth corner.
Skin on anteroventral surface of lower jaw cor-
rugated and papillate. Palate behind heavily
fringed maxillary valve with three strong short
ridges, a medial longitudinal ridge and a pair of
diagonal lateral ridges; two short ridges also pres-
ent behind the three palatine ridges. Floor of
mouth with a transverse row of three elongated
oral papillae, a larger medial papilla and a small-
er lateral one behind each end of dental band; a
pair of additional large papillae just behind and
lateral to medial papilla. Upper jaw with 5-7 and
lower jaw with 7-9 functional tooth series. Teeth
in quincunx pavement, close-set, with crowns
adjacent to one another. Teeth similar in upper
and lower jaws, uppers slightly larger than low-
ers; teeth varying continuously lateral to sym-
physis, largest and longest relative to width at or
near symphysis, smallest and shortest near mouth
corners. Upper dental band with a small knob
of slightly enlarged teeth at symphysis, separated
from similar weak knobs on either side by
depressions with smaller teeth; lower dental band
with a corresponding symphyseal depression into
which upper symphyseal knob fits, and a pair of
very low lateral knobs which fit into upper par-
asymphyseal depressions. Teeth of both jaws with
moderately high (females) to very high (males),
peaked, broad-tipped (females) or acutely point-
ed, cuspidate (males) crowns shaped like mush-
room caps, with a strong, sharp cutting edge
(males) or a broad, blunt carina or transverse
keel (females), a strong basal ledge, and a con-
cave, inset basal groove separating crown from
root. Root small, moderately high, pedicellate,
bilobed basally, with a transverse groove and
nutrient foramen. Dentition sexually dimorphic;
teeth of males with triangular, concave, cuspi-
date crowns; females with low, rhomboidal,
truncated, cuspless crowns.

Dorsal surface of disc either completely naked,
without denticles (2 fetuses, the holotype, and a

294

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18

FIGURE 5. Dasyatis garouaensis, 342-mm mature female, Cross River near Mamfe, Cameroun (CAS 49147).

340-mm adult male), or with a variable number
of small, heart-shaped or circular denticles bur-
ied in skin on middorsal area (remainder of spec-
imens examined). Two large specimens (includ-
ing 342-mm male from Mamfe) have a few
scattered denticles along midline of back; two
others have a small rectangular area of flat den-
ticles centered on midscapular region but not
extending outwards to cover middle third of disc
as in D. ukpam, D. margarita, and D. marga-
ritella. Five large specimens had 3 or 4 large,
flattened, wedge-shaped denticles in midscapular
region, but these not formed as domed, rounded
pearl spines (although probably homologous to
pearl spines).

Neurocranium, examined from radiographs,
generally similar to that of Himantura signifer
as described in Compagno and Roberts (1982),
except frontoparietal fontanelle shorter and
broader, and posterior margin of nasal capsules
more transverse; width of nasal capsules about
75% of nasobasal cranial length.

Pelvic girdle (Fig. lOc-d) less arched than that
of D. ukpam, convex anteriorly and without a
medial prepubic process or a median angle. Pel-

vic girdle of D. garouaensis differs from that of
D. ukpam as well as D. margarita and D. mar-
garitella in having large, laterally expanded lat-
eral prepelvic processes; these were prominent
on all D. garouaensis radiographed, including
specimens from Mamfe, Lagos, and Benue River
in Nigeria and Cameroun. Pelvic girdle also with
short ischial processes, long, slender iliac pro-
cesses, and 4 obturator foramina.

Claspers of adult male short and stout, length
of outer margin 10.2-1 1.6% of disc width, oval
in cross section and somewhat depressed; height
about 3/5 of width at midlength. Dorsal surface of
clasper slightly flattened, ventral surface broadly
convex, lateral edge convexly arched, medial edge
undulated, and tip bluntly pointed. Apopyle on
anterodorsal surface, connected to hypopyle by
an open, posteriorly curved clasper groove.
Clasper glans simple, dorsal lobe supported by
dorsal marginal and terminal cartilages, ventral
lobe supported by ventral marginal and terminal
cartilages as well as ventral covering piece. No
structures inside hypopyle. A large pseudopera
laterally situated on ventral lobe below hypopyle;
a small pseudosiphon on dorsomedial surface of

COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS

295

dorsal lobe, its cavity lying entirely below flange
of dorsal marginal cartilage. Ventral lobe without
clasper spur or associated terminal (T3 cartilage).
Clasper skeleton simple, with two cylindrical
basal segments connecting pelvic basipterygium
to axial cartilage. Axial cartilage cylindrical,
nearly straight, with rodlike tip reaching ends of
terminal cartilages. Beta cartilage present, a long,
slender, separate, flattened plate running along
lateral surface of clasper skeleton from posterior
end of basipterygium to anterior end of dorsal
marginal cartilage. Dorsal marginal cartilage
broad and subquadratetriangular, with a diago-
nally truncate posterior edge, broad medial flange,
and narrow lateral flange that forms roof of clasp-
er groove. Ventral marginal cartilage a narrow,
laterally expanded plate on axial cartilage, with
a straight lateral margin forming floor of clasper
groove. Dorsal terminal cartilage large, broad,
wedge-shaped, and axially convex, with a broad
anterior base articulating with postero ventral edge
of dorsal terminal, a narrow posterior tip op-
posite tip of axial cartilage, and medial edge ar-
ticulating with axial cartilage. Ventral terminal
cartilage large, complex, oval, and scoop-shaped,
with a broad, arched, lateral flange forming roof
of pseudopera and a recurved, expanded pos-
teroventral tip forming a partial floor for it along
with ventral covering piece. Ventral covering
piece large, elongate-oval, broadly convex ven-
trally, and scoop-shaped, enclosing terminal car-
tilages and rear tips of marginals ventrally.

Dasyatis ukpam (Smith, 1863)

(Figures 6-9)

Hemitrygon ukpam SMITH, 1863:69 (type-locality Old Calabar
River, Nigeria); FOWLER, 1 936: 1 26 (placed in synonymy of
D. centroura); FOWLER, 1969:186 (in synonymy of D. cen-
trourd).

Trygon ukpam GUNTHER, 1870:480 (description after Smith,
1863, placed in Trygon = Dasyatis).

Dasyatis margarita LOUBENS, 1 964: 1 1 (freshwater lakes south
of Lambarene district, Ogooue basin; presumably no spec-
imens preserved).

Dasyatis ukpam STEHMANN, 1981:4 (in key to marine Dasyatis
of West Africa); COMPAGNO AND ROBERTS, 1982:321 (ref-
erence).

MATERIAL EXAMINED. — BMNH 1874.5.23.1, syntype, 266-
mm late fetal male, Old Calabar River, Nigeria; USNM 219780,
520-mm immature female, Lake Ezanga, Ogooud River sys-
tem, Gabon; CAS 42761, 650-mm immature female, Lake
Ezanga near Nzame-Akesile village, Ogooue River system, Ga-
bon; MNHN 1979-244, 499-mm immature female, Booue,
Ogoou6 River, Gabon; MRAC 55778, 361 -mm immature fe-
male, Binda, Congo (Zaire) River, Zaire.

DIAGNOSIS.— D. ukpam is a very large and

thick-bodied freshwater dasyatid, probably
growing much bigger than our largest specimen,
a 650-mm immature female. Newborn young,
266 mm, are as large as the largest known D.
margaritella and far larger than newborn young
of D. margarita and D. garouaensis. Entire dor-
sal surface of disc covered with stout-spined den-
ticles (smooth in newborn) (peripheral portions
of disc without denticles in all other West African
dasyatids except the very large marine species D.
centroura, with a diamond-shaped disc, and Uro-
gymnus, with an extremely thick disc). Sting
greatly reduced in size or absent (sting invariably
absent in Urogymnus but normally present and
relatively large in all other West African dasy-
atids).

Disc oval, very thick. Medial lobe narrow-
based and short, its anterior margin broadly con-
vex; disc depth 13.3-15.7% of disc width, disc
width 0.9-1.0 times disc length; dorsal surface
of disc entirely covered with denticles at all free-
living stages except possibly newborn; small,
heart-shaped or circular denticles covering mid-
dle of disc, small prickles on sides of disc, with
scattered large, conical, stellate, sharp denticles
(absent in newborn and fewer in small immatures
than large) and usually 1-3 small to moderate-
sized midscapular pearl spines on disc; no sting,
or a single small, possibly vestigial one, only 6.5%
of disc width in 520-mm specimen; eyes small
and strongly elevated, eyeball lengths 2.1-3.3
times in interorbital space; interorbital space 1.5-
1 .9 times in preorbital length; spiracles large and
high; floor of mouth with 4 or 5 elongated pa-
pillae; total tooth rows 38-46/38-48; pelvic fins
short, anterior margins 16-19% of disc width;
tail uniformly tapering to a slender whiplash, its
length from vent when intact 2.0-2.9 times disc
width; base of tail circular in section; a very low
ventral tailfold but no dorsal fold or keel; disc
and pelvic fins dark brown or gray-brown above,
without spots or prominent markings, white be-
low except for broad dark margins; tail blackish
except for underside of its base; intestinal valve
turns 19-20; total pectoral radials 142-148, total
vertebral centra 108-1 22 and total segments 155
(in one specimen), vertebral centra ending in front
of sting or extending to its origin.

Proportional measurements (as percent of disc
width) and counts are given in Table 1 . Preoral
length 2.3-2.6 times internarial width and 0.9-
1 .0 times width between first gill openings. Snout
broad, angle in front of eyes 123-132°. Spiracle

296

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18

FIGURE 6. Dasyatis ukpam, 266-mm fetal male syntype, Old Calabar, Nigeria (BMNH 1874.5.23:1).

FIGURE 7. Dasyatis ukpam, 650-mm immature female, Lake Ezanga, Ogooue basin, Gabon (CAS 42761).

COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS

297

FIGURE 8. Dasyatis ukpam, 520-mm immature female, Lake Ezanga, Ogoou6 basin, Gabon (USNM 219780).

length 1.5-2.0 times eyeball length, 0.8-1.0 times
internarial width, and 1.8-2.2 times in distance
between fifth gill openings. Internal gill openings
with close-set transverse ridges on gill arches.
Nasal curtain with a fringed, slightly concave or

trilobate posterior margin. Mouth weakly arched,
midline of lower jaw slightly indented; a shallow
to deep, curved groove extending posteriorly from
posterior nasal flap around corners of mouth.
Skin on ventral surface of lower jaw more or less

FIGURE 9. Dasyatis ukpam, 361-mm immature female, lower Zaire River at Binda, Zaire (MRAC 55778).

298

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18

FIGURE 1 0. Pelvic girdle (dorsal view) in West African Das-
yatis: (a) D. ukpam, 266-mm fetal male; (b) D. ukpam, 361-
mm immature female; (c) D. garouaensis, 342-mm mature
female; (d) D. garouaensis, 255-mm adult male; (e) D. mar-
garita, 200-mm immature female; (0 D. margaritella, 190-
mm mature male. Note prominent lateral prepelvic processes
in D. garouaensis. Scale bar = 2 cm.

corrugated and papillate. Palate behind fringed
maxillary valve with three strong, short ridges,
a medial longitudinal ridge and a pair of diagonal
lateral ridges. Two pairs of long, low ridges be-
hind the three palatine ridges. Floor of mouth
with a transverse row of usually three moderately
large, elongated oral papillae, a medial papilla
(absent in syntype) and a lateral papilla behind
each end of dental band; a pair of large papillae
just behind and lateral to medial papilla.

Upper jaw with about 4 and lower jaw with 6
functional tooth series. Teeth in quincunx pave-
ment, close-set, with crowns closely adjacent to
one another. Teeth similar in upper and lower
jaws, uppers slightly larger than lowers at sym-
physis, varying continuously lateral to symphy-
sis, teeth largest and longest relative to width at
or near symphysis and smallest and shortest at
mouth corners. Upper dental band with a low
knob of slightly enlarged teeth at symphysis, sep-
arated from similar knobs on either side by
depressions with smaller teeth; lower dental band
with a weak symphyseal depression into which
symphyseal knob of upper jaw fits; a pair of low
lateral knobs fit into depressions in upper dental
band. Teeth of both jaws in females with mod-
erately high, broad-tipped, cuspless crowns
shaped like mushroom caps, with a transverse,
blunt keel or carina, strong transverse ridges on
both labial and lingual sides, and a strong basal
ledge and concave, inset basal groove separating
crown and root. Roots moderately large, high,
pedicellate, bilobed basally, with a transverse
groove and nutrient foramen. Teeth of adult males

unknown, but probably differ from those of fe-
males.

Dorsal surface of 266-mm late fetus covered
with small flat denticles on middle third of disc
and base of tail to its abbreviated sting, including
dorsal surface of cranium, branchial region, scap-
ular region, and abdominal area; in addition,
specimen has two small, slightly elevated, en-
larged rounded denticles or pearl spines in medi-
scapular area. The larger, free-living specimens
examined have outer two-thirds of disc, snout,
and tail posterior to sting region with small to
moderately large, conical, prickle-like denticles
in addition to flattened denticles covering mid-
belt; they also have massive, conical, erect, flut-
ed, sharp denticles or thorns over much of dorsal
surface of disc and tail base. Large thorns fewest
on smallest (36 1-mm) specimen, most numerous
on largest (650-mm), suggesting that they be-
come more numerous with growth. These thorns
make dorsal surfaces of large D. ukpam ex-
tremely rough, and, as noted by Smith (1863),
difficult to handle. Free-living specimens ex-
amined have one or two round, enlarged pearl
spines on mediscapular region.

Neurocranium observed on radiographs but it
and other skeletal parts obscured by thickness of
disc and heavy covering of denticles in this
species. Cranium apparently similar to that of
Himantura signifer as described in Compagno
and Roberts ( 1982), but with a straighter anterior
margin to its nasal capsules.

Pelvic girdle (Fig. lOa-b) broadly arched,
semicircular, relatively thick, with a medial an-
terior angle but no medial prepubic process. Lat-
eral prepubic processes low, rounded, and lobate;
iliac processes well-developed; ischial processes
short. Five obturator foramina present on one
specimen.

Claspers of mature male not available.

DISCUSSION

It has been known for some time that stingrays
identified as Dasyatis margarita represent two
species (Daget and Iltis 1965; Blache et al. 1970;
Stehmann 198 1). It appears that the first ichthy-
ologist to become aware of this was the late J.
Cadenat; he recognized that the two species differ
greatly in weight, the one not exceeding 1 kg
while the other attains easily 1 5-20 kg and per-
haps much more (quoted in Daget and Iltis 1 965:
15). The small species is D. margaritella, the

COMPAGNO AND ROBERTS: WEST AFRICAN STINGRAYS

299

large one D. margarita. Both are common and
widely distributed in shallow water along the coast
of West Africa. Most accounts of D. margarita
are based on D. margaritella or on both D. mar-
garitella and D. margarita. D. margaritella is
more common than D. margarita in museum
collections and perhaps also in nature. We are
unaware of any difference in habitat preference.

It is remarkable that such notable animals as
freshwater stingrays remain so poorly known.
We believe that they will be found in additional
river basins in West Africa and that possibly
additional freshwater species are present. We have
heard that stingrays occur in the Sanaga basin in
Cameroun, particularly in Lac Ossa, but have no
material evidence for this. It is curious that no
Dasyatidae have been reported from rivers west
of Nigeria. This might be due to insufficient col-
lecting.

A stingray was reported from the Cross River
at Mamfe Pool by Sanderson (1937), but the
specimen was not preserved, and the account is
so extraordinary that we hardly know what to
make of it. According to Sanderson, the ray was
"diamond-shaped, like all fish of this class, and
measured from the tip of one lateral point to the
tip of the other, four feet eight inches; from the
snout to the base of the tail, five feet eleven inches;
and from the base to the tip of the tail, which
had no fin, five feet two inches. Emerging from
the upper edge of the tapering whip-like tail near
its base was a long, straight, sharp spine or sting,
one foot seven inches in length." He went on to
say that the arrival of this monster altogether un-
hinged his sense of logic, so perhaps it also af-
fected his ability to observe and record accu-
rately. "That it was still alive and therefore
undoubtedly caught in Mamfe Pool, as the na-
tives stated, was almost incredible, because this
bit of water was nearly three hundred miles from
the sea. I therefore had to adjust myself to the
idea that such things are true fresh-water animals
indigenous to the great rivers of Africa. Why do
not natural history books depict these fish in-
stead of the everlasting crocodile?" Why not, in-
deed? Sanderson provides as good an answer as
any: "We didn't want the brute because we were
not collecting fish, but we photographed him
alongside sundry natives and inanimate objects
and purchased the sting." Unfortunately the pho-
tograph was not published and the sting had to
be discarded: "When this sting got really dry it

split longitudinally and opened like a star, re-
vealing a clear crystalline plug within. This sub-
stance gradually broke up under the damp at-
mospheric conditions; some of it dropped in water
fizzed furiously. I could not find anything that
would preserve it among our selection of trav-
elling drugs and chemicals."

We have seen dried stings of various rays with-
out observing anything like the disintegration re-
ported by Sanderson, and suspect he had a few
chuckles and perhaps something else while con-
cocting this giant sting. The fishermen at Mamfe,
who regularly catch D. garouaensis, maintained
that it is the only species of ray in the Cross River
from Mamfe to the Nigerian border and that the
example caught and preserved during the junior
author's visit was nearly as large as the largest
they had ever caught. When shown a photo-
graphic print of D. ukpam they indicated that
this species was unknown to them. Concerning
sting size, the largest sting we have observed on
a West African freshwater ray is under four inches.
In Dasyatis centroura, which probably has the
largest sting of any marine species in West Africa,
the sting of a specimen with a disc width of four
feet eight inches would be only about six or seven
inches. We conclude that D. garouaensis is the
only species of stingray in the Cross River in the
vicinity of Mamfe.

Identification of a specimen of D. garouaensis
from Lagos may indicate that the species occurs
in Lagos Lagoon and in the rivers flowing into
it. Unfortunately the specimen is not accom-
panied by information on habitat, and we cannot
rule out the possibility that it was caught in the
Niger River and transported to Lagos.

Although the title of the paper in which D.
ukpam was described states that it lives in the
Old Calabar River, and the text indicates that it
lives as much as 1 50 miles upriver (Smith 1863),
this is based on hearsay. It should be noted that
the vernacular name "ukpam" or "okpam" is a
generical term for stingrays. At Mamfe this name
is employed by present-day speakers of the Man-
yu and Ejagam languages for D. garouaensis.

The occurrence of an unidentified stingray in
the Ogoou6 basin was first brought to our atten-
tion by an ichthyological colleague, Dr. Jaques
Gery, who related to the junior author that he
had observed two rays in the Ivindo near Ma-
kokou while collecting characins and other small
fishes in 1964. The Ivindo flows into the Ogoou6

300

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 18

near Booue, where one of our specimens of D.
ukpam was collected, but there are some for-
midable waterfalls on the Ivindo below Mako-
kou and the species there might be different. The
Zaire locality for D. ukpam, Binda, is on a rel-
atively narrow and swift-flowing portion of the
lower Zaire (Congo) River about 100 km upriver
from the mouth of the river at Banana and 35
km downriver from the end of the mainstream
rapids of the lower Zaire River near Matadi. No
stingrays are known from the interior of the Zaire
or Congo basin. So far as we have been able to
determine D. garouaensis and D. ukpam are the
only living freshwater stingrays known from Af-
rica.

ACKNOWLEDGMENTS

For providing information and facilitating ex-
amination of specimens we wish to thank Al-
wyne Wheeler, Mandy Holloway, and Oliver
Crimmen, British Museum (Natural History); D.
F. Thys van den Audenaerde, Musee Royale de
1'Afrique Centrale, Tervuren, Belgium; and
Thomas B. Thorson, School of Life Sciences,
University of Nebraska. Fieldwork in Gabon was
facilitated by the Centre National pour la Re-
cherche Technique et Scientifique and particu-
larly M. le Commissaire Nzoghe-Ngueme. We
thank Jeanne Byloghe and Joseph Mebiaghe of
Tame and Nzame-Akesile villages for catching
Dasyatis ukpam, and Pierre Ville of Geri Con-
sult, who also obtained a specimen of this species
for us. Permission for fieldwork in Cameroun
was kindly granted by the Office National pour
la Recherche Scientifique et Technique. At
Mamfe the junior author was assisted by S. T.
Mbianyor of the Forestry Department, John
Corrao and Alan Ferguson of the Peace Corps,
and a number of excellent local fishermen.

LITERATURE CITED

BIGELOW, H. B., AND W. C. SCHROEDER. 1953. Fishesofthe
western North Atlantic. Sears Found. Mar. Res., Mem. No.
1(2).

BLACHE, J., J. CADENAT, AND A. STAUCH. 1970. C16s de de-

termination des poissons de mer signa!6s dans 1'Atlantique

oriental. Faune Trop. (ORSTOM, Paris), No. 18.
CASTELLO, H. 1973. Sobre la correcta posicion sistematica

de la raya de agua dulce africana (Chondrichthyes, Dasy-

atidae) (Republic Federal del Camerun). Trab. V Congr. La-

tinoamer., Zool. 1:67-71.
CASTEX, M. 1967. Fresh water venomous rays. Pages 167-

176 in Animal toxins. Pergamon Press, Oxford and New

York.
COMPAGNO, L. J. V., AND T. R. ROBERTS. 1982. Freshwater

stingrays (Dasyatidae) of Southeast Asia and New Guinea,

with description of a new species of Himantura and reports

of unidentified species. Environ. Biol. Fishes 7(4):32 1-339,

12 figs.
DAGET, J., AND A. ILTIS. 1965. Poissons de Cote d'lvoire

(eaux douces et saumatres). M6m. Inst. franc. Afr. noire no.

74.
DAGET, J., AND A. STAUCH. 1963. Poissons de la partie Ca-

merounaise du bassin de la Benou6. Mem. Inst. franc. Af-

rique noire 68:85-107.
FOWLER, H.W. 1936. The marine fishes of West Africa. Bull.

Amer. Mus. Nat. Hist. 70(1): 1-606.
. 1969. A catalog of world fishes (XI). Quart. J.Taiwan

Mus. 22(3-4): 125- 190.
GUNTHER, A. W. 1870. Catalogue of the fishes in the British

Museum. 8.
KREFFT, G. 1968. Knorpelfische (Chondrichthyes) aus dem

tropischen Ostatlantik. Atlantide Rep. 10:33-76, pis. 3-6.
LOUBENS, G. 1964. Travaux en vue de developpement de la

peche dans le bassin inferieurde 1'Ogooue. Publ. Cent. Tech.

Forest. Trop., Nogent-sur-Marne (Seine) 27:1-51.
RAFINESQUE, C. S. 1810. Caratteri di alcuni nuovi generi e

nuove specie di animali (principalmente di pesci) e piante

della Sicilia, con varie osservazioni sopra i medisimi. Pa-
lermo, 105 pp, 20 pis.
REID, G. M., AND H. SYDENHAM. 1979. A checklist of lower

Benue River fishes and an ichthyogeographical review of the

Benue River (West Africa).
SANDERSON, I. 1937. Animal treasure. Viking Press, New

York, 325 pp.
SMITH, J. A. 1863. Notice of the ukpam, a large species

(probably new) of sting ray ( Trygon, Cuvier), found in the

Old Calabar River, Africa. Proc. Roy. Phys. Soc. Edinburgh

1859-62,2:64-69.
STAUCH, A., AND M. BLANC. 1962. Description d'un selacien

rajiforme des eaux douces du Nord-Cameroun, Potamotry-

gon garouaensis n. sp. Bull. Mus. Nat. Hist. Natr. (Paris)

34(2): 166-1 71.
STEHMANN, M. 1981. Dasyatidae. In Fischer, W., G. Bianchi,

and W. B. Scott (eds.). FAO species identification sheets for

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THORSON, T. B., AND D. E. WATSON. 1975. Reassignment of

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grounds. Copeia 1975(4):701-712, figs. 1-3.

CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, California 94118

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 19, pp. 301-315, 12 figs., 1 table. December 11, 1984

REVISION OF EASTERN PACIFIC CATALUFAS

(PISCES: PRIACANTHIDAE) WITH DESCRIPTION OF A NEW

GENUS AND DISCUSSION OF THE FOSSIL RECORD

John E. Fitch

Research Associate, California Academy of Sciences,
San Francisco, California 94118

and
Stephen J. Crooke

California Department of Fish and Game,
Long Beach, California 90802

ABSTRACT: Four species of catalufas inhabit eastern Pacific waters: Cookeolus hoops (Schneider, 1801),
Heteropriacanthus cruentatus (Lacepede, 1801), Pseudopriacanthus serrula (Gilbert, 1891), and Priacanthus
alalaua Jordan and Evermann, 1904. Each of these species is illustrated, and diagnostic characters, meristic
data, morphometric measurements, maximum size, geographic range, depth distribution and other informa-
tion also are presented. Heteropriacanthus is a new generic name for Priacanthus cruentatus, a cosmopolitan
species that differs in numerous salient features from the species assignable to Priacanthus (i.e., alalaua,
arenatus, hamrur, macracanthus, meeki and tayenus). An identification key is presented. Otoliths (sagittae)
and scales of the four eastern Pacific species also are illustrated. The only reported priacanthid fossils are
from the Eocene of Europe. Of the six species, Pristigenys substriata is known from skeletal remains and is
unquestionably a priacanthid. Only two of the five species described from otoliths, Pristigenys bella and P.
dentifer, appear to be priacanthids, but there is no assurance they can be assigned to Pristigenys, since none
of the skeletal "imprints" of P. substriata contained otoliths, nor do all of the otoliths assigned to these two
species appear to be correctly identified.

INTRODUCTION partment of Fish and Game (DFG) biologists,
For years, fishermen aboard long-range sport- skippers and crew members of these vessels corn-
fishing boats that operated out of San Diego menced fishing for and saving miscellaneous
sought only such large game species as yellowfin small fishes caught at the Revillagigedo Islands,
and bigeye tuna (Thunnus albacares and T. obe- Alijos Rocks, and other fishing spots off southern
stis), wahoo (Acanthocybium solanderi), yellow- Baja California.

tail (Seriola lalandi), giant sea bass (Stereolepis Among the first of these incidentally caught

gigas) and several kinds of large serranids (Epi- species turned over to DFG personnel were a

nephelus spp. and Mycteroperca spp.). During half-dozen catalufas that appeared to represent

1978, however, at the urging of California De- three species of Priacanthus. A literature search

[301]

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19

FIGURE 1 . Type specimen ofAnthias macrophthalmus Bloch, 1 792 (ZMB 8 1 56). Photo courtesy of Hans-J. Paepke, Museum
fur Naturkunde der Humboldt-Universitat zu Berlin.

and examination of all the priacanthids in the
fish collections at Scripps Institution of Ocean-
ography (SIO), Natural History Museum of Los
Angeles County (LACM), University of Califor-
nia, Los Angeles (UCLA) and California Acad-
emy of Sciences (CAS) revealed that, until then
(Dec. 1978), any priacanthid collected in the
eastern Pacific that was not judged to be either
Pseudopriacanthus or Cookeolus, automatically
had been relegated to Priacanthus cruentatus (see
also Fitch and Schultz 1978).

Once it became obvious that we were dealing
with several species, the differences in scale
counts, pelvic fin pigmentation, eye diameter,
otoliths, and gas bladder morphology became
equally obvious. During the succeeding four years,
in an effort to determine exactly what species we
were observing, we examined several hundred
priacanthids from throughout the world. As a
result, we arrived at the conclusion that there
were four species of catalufas in the eastern Pa-
cific: Cookeolus hoops, Priacanthus alalaua,
Priacanthus cruentatus and Pseudopriacanthus
serrula.

In the course of our investigation, we found

that otoliths (sagittae) and gas bladders of Pria-
canthus alalaua were so radically different from
otoliths and gas bladders of P. cruentatus that a
new generic name was needed for one of the two.
This necessitated determining the condition of
the otoliths and gas bladder of Bloch's (1792)
Anthias macrophthalmus, since it was the type-
species for Priacanthus Oken, 1817. Fortunately,
Bloch's type specimen, a skin from the right side
of the fish, still exists in the Museum fur Natur-
kunde der Humboldt-Universita't zu Berlin
(ZMB8156), and Hans-J. Paepke, Curator of
Fishes, sent us an excellent photograph of this
specimen (Fig. 1). Wayne Starnes (pers. comm.),
to whom we sent a copy of the photograph, has
confirmed that Bloch's Anthias macrophthalmus
is conspecific with Sciaena hamrur Forsskal,
1775.

KEY TO EASTERN PACIFIC PRIACANTHIDAE

1 a. Pored scales in lateral line 3 5 to 40; dorsal
soft rays 1 1 (rarely 10 or 12); anal rays 10;
dorsal profile turns abruptly downward
under base of soft portion of second dorsal
fin; ventral profile turns abruptly upward

FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS

303

FIGURE 2. Pseudopriacanthus serrula. Photo by Jack W. Schott.

under anal fin base; ctenii on scales in par-
allel species, presenting a corrugated or

waffled appearance _

Pseudopriaca nth us serrula

Ib. More than 50 pored scales in lateral line;
1 2 (rarely) or more dorsal soft rays; 1 2 or
more anal rays; dorsal and ventral profiles
tapering gradually to caudal peduncle;
ctenii on scales not as above 2

2a. Eighteen to 20 scale rows between 8th dor-
sal spine and lateral line; more than 20
rows on dorsum of caudal peduncle; anal
rays 13; peduncle depth about equal to
horizontal eye diameter; dorsal, anal and
caudal fins yellow, edged with black; ctenii

thick and bristlelike, in rows

Cookeolus boops

2b. Eight to 10 scale rows between 8th dorsal
spine and lateral line; fewer than 1 5 scale
rows on dorsum of caudal peduncle; anal
rays 14; peduncle depth fits about 1 '/4 times
into horizontal eye diameter; dorsal, anal

and caudal fins never with yellow or tipped
with black; ctenii thin and filamentous ..... 3

3a. Second dorsal, anal and caudal fin mem-
branes red, without spotting; pelvic fin
membranes jet black; gas bladder with an-
teriorly projecting "ears" that extend to
the otic bullae, and posteriorly projecting
horns that reach to above end of anal fin;
otoliths (sagittae) with a ventral keellike
blade and centrally positioned pronglike
rostrum; ctenii filamentous, in rows; pre-

opercle completely scaled

Priacanthus alalaua

3b. Second dorsal, anal and caudal fin mem-
branes with rust-colored spotting (dusky
in formalin-preserved specimens); pelvic
fin membranes lightly pigmented to clear;
gas bladder contained entirely within body
cavity, without ears and no anterior or
posterior projections; otoliths (sagittae)
lack keellike blades, oval in outline, with
normal, anteroventral rostrum; ctenii fil-

304

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19

amentous, only on scale margin, sparse;
posterior margin of preopercle without

scales _...

Heteropriacanthus cruentatus new genus

SPECIES ACCOUNTS
Pseudopriacanthus serrula (Gilbert, 1891)

(Figure 2)

DIAGNOSTIC CHARACTERS. — 35 to 40 pored
scales in lateral line; 1 0 to 1 2 (typically 1 1 ) dorsal
soft rays; 10 anal rays; ctenii on scales in a par-
allel series (Fig. 3d) presenting a corrugated or
waffled appearance; dorsal profile turns abruptly
downward, becoming nearly vertical, under sec-
ond dorsal fin base; ventral profile becomes near-
ly vertical under anal fin base.

MERISTIC DATA.-D. X.10-12; A. 111,10; P.
17; GR 6-7 + 15-18 = 21-25; pored lateral line
scales 35-40; vertebrae 10 + 13 = 23. Twenty-
three of the 25 specimens we examined for me-
ristic data had 1 1 dorsal soft rays, one had 10,
and one had 12.

MAXIMUM SIZE.— The largest individual we
observed was a female 274 mm SL (353 mm TL)
that weighed 1 300 g. This was one of 24 indi-
viduals caught in gill nets in "30-40 fm" (55-
73 m) offMagadalena Bay, Baja California, dur-
ing March 1976.

RANGE.— Monterey Bay, California, to Talara,
Peru (Fitch and Lavenberg 1975) and at most
offshore islands from the Coronados to the Ga-
lapagos in 3.6 to more than 100 m (DFG, un-
published data).

REMARKS. — Morphometric data were taken on
only 1 3 of the more than 60 individuals we ex-
amined (Table 1). The smallest of these (34 mm
SL, LACM 22796) had a relatively short pelvic
fin (41% of SL) as compared with the Eocene
fossil Pristigenys substriata (Fig. 1 1) and pelagic
stages of the extant Cookeolus hoops (Fig. 5, Ta-
ble 1). Fritzsche and Johnson (1981) considered
Pseudopriacanthus a junior synonym of Pristi-
genys, but for reasons given later, we believe both
genera are distinct and valid.

Of perhaps 20 individuals that have been
caught or observed in Californian waters, one
each was from Monterey Bay and off San Luis
Obispo, the rest have been from south of Pt.
Dume. Scuba divers report that P. serrula is un-
afraid and can be picked up by hand when en-
countered in its natural surroundings.

Cookeolus hoops (Schneider, 1801)
(Figure 4)

DIAGNOSTIC CHARACTERS. — 1 8 to 20 scale rows
between eighth dorsal spine and lateral line; more
than 20 rows of scales on dorsum of caudal pe-
duncle; ctenii on scales thick and bristlelike (Fig.
3a), in rows; anal rays 12-13; dorsal, anal and
caudal fins yellow, edged with black; peduncle
depth about equal to horizontal eye diameter;
pelvic fins of pelagic juveniles comprise 50-70%
of SL or more.

MERISTIC DATA.-D. X,13; A. 111,12-13; P.
18-19; GR 6-8 + 17-18 = 23-26; pored lateral
line scales 53-61; vertebrae 10 + 13 = 23. One
of the 35 specimens we examined for meristic
data had 12 anal rays, the rest had 13.

MAXIMUM SIZE.— The largest individual we
observed was a female 397 mm SL (507 mm TL)
that weighed 2725 g. It was caught by a sport-
fisherman at Alijos Rocks, Baja California, Mex-
ico, in 1974 in "18 fm" (33 m) of water. This
fish (LACM 34253) was erroneously identified
by Fitch and Schultz (1978) as Priacanthus
cruentatus. In the western North Atlantic, a C.
boops has been reported that was 507 mm SL
and weighed 5.2 kg (Anderson et al. 1972).

RANGE.— Worldwide in tropical and subtrop-
ical seas; in the eastern Pacific from Alijos Rocks,
Mexico (24°57'N, 115°45'W) to 10°N 98'W
(LACM 30506-1). Although adults have been
caught in water as shallow as 30 m in the eastern
Pacific, they are most commonly hooked at depths
"exceeding 40 fm" (73 m) and have been taken
as deep as "75 fm" (137 m). Pelagic juveniles
have been captured at the surface at scattered
offshore localities between the Tres Marias Is-
lands and the Gulf of Tehuantepec. In the west-
ern North Atlantic, C. boops has been taken at
depths exceeding 365 m (Anderson et al. 1972).

REMARKS.— Although Cookeolus was not rec-
ognized from the eastern Pacific until Fritzsche
(1978) reported upon six pelagic juveniles rang-
ing from 148 to 226 mm SL, adults had been
taken at Alijos Rocks as early as 1970, but were
erroneously identified as Priacanthus cruentatus
(Fitch and Schultz 1978). Not until 1978, when
long-range sportfishing boats started bringing in
fair numbers of adult C. boops from Alijos Rocks,
the Revillagigedo Islands and Hurricane Bank
(16°52'N, 117°28'W) were they recognized for
what they were. Most of the confusion in iden-
tification had resulted from the relatively shorter

FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS

305

FIGURE 3. Scales of eastern Pacific Priacanthidae (scale height and length, in mm, in parens): a) Cookeolus boops (4.6 by
3.5); b) Priacanthus alalaua (3. 1 by 3. 1); c) Heteropriacanthus cruentatus (1 .9 by 2.4); d) Pseudopriacanthus serrula (3.6 by 4.0).
SEM photos by Richard Huddleston.

pelvic fins of the bottom-dwelling adults and the
lack of spots and blotches which are so typical
of the pelagic juveniles (Fritzsche 1978).

Although pelvic fin length commences to
shorten when Cookeolus takes up a bottom-
dwelling existence, six of the specimens we used

in obtaining morphometric data (Table 1) still
had pelvic fins that exceeded 40% of SL. All six
of these fish, ranging from 197 to 267 mm SL
(Fig. 5), had been hooked on the bottom. On the
other hand, two bottom-dwelling specimens (222
and 257 mm SL) had pelvic fins that had short-

306

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19

FIGURE 4. Cookeolus hoops. Photo by Lee Stockland.

PERCENT OF
STANDARD LENGTH

PELVIC FIN LENGTH

•STANDARD LENGTH(MM)

160 180 200 220 240 260 280 300 320 340 360 38O 400

FIGURE 5. Pelvic fin length (as percent of SL) plotted against SL for 32 specimens of Cookeolus boops.

FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS

307

FIGURE 6. Juvenile Cookeolus boops, 211 mm SL, taken on hook and line in "15 fms" at Alijos Rocks. Photo by Paul
Gregory.

TABLE 1 . COMPARATIVE MEASUREMENTS FOR THE FOUR EASTERN PACIFIC PRIACANTHIDS (in percent standard length).

Species and number of specimens measured

Measurement

Pseudopriacanthus
serrula

Cookeolus
boops

Priacanthus
alalaua

Heteropriacanthus
cruentatus

Number of specimens

Standard length (mm)

34-261

151-392

215-261

81.5-245

Head length

36.1-41.2

31.2-37.4

29.4-33.3

28.6-36.4

Horizontal eye diameter

14.3-18.0

9.6-13.9

12.5-14.1

10.2-15.2

Snout length

9.0-11.4

9.8-11.7

9.5-10.5

9.0-10.1

Maxillary length

20.0-21.6

16.6-19.2

14.7-16.1

15.5-17.1

Bony interorbital width

7.9-10.2

8.2-10.5

8.0-9.1

8.4-10.1

Snout to 1st dorsal fin insertion

33.7-36.4

28.0-33.3

30.0-32.3

29.7-32.3

Snout to pectoral fin insertion

37.6-40.8

31.7-37.9

30.9-34.8

30.1-32.9

Snout to pelvic fin insertion

44.8-46.8

34.0-44.0

33.3-38.8

35.4-38.3

Snout to anal fin insertion

70.9-75.6

58.1-67.9

55.3-60.9

51.3-59.0

Dorsal fin insertion to pelvic insert

48.4-53.6

37.7-48.8

36.4-39.1

35.0-40.3

Depth perpendicular to AS,

48.8-54.2

37.1-49.2

35.6-38.6

34.3-39.8

Caudal peduncle depth

12.8-14.0

9.6-12.3

7.7-8.2

8.3-10.5

Pectoral fin length

19.3-23.2

18.9-24.8

20.8-23.2

17.0-21.0

Pelvic fin length

25.6-35.4'

27.0-70.22

25.3-31.4

19.7-26.7

Longest gill raker

5.9-8.8

5.0-7.2

4.1-5.4

4.4-5.2

1 Pelvic fin length of 34 mm specimen (41.2%) not included, remaining specimens 172 to 261 mm SL.

2 See Figure 5.

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19

-•

•

• • *

•
.

•

FIGURE 7. Priacanthus alalaua. Photo by Lee Stockland.

ened to 34% of SL (Figs. 5 and 6). The largest
pelagic juvenile reported by Fritzsche (1 978) was
226 mm SL. Based upon these lengths, in the
eastern Pacific, Cookeolus remains in the pelagic
environment until it reaches a size of approxi-
mately 200-250 mm SL.

Interestingly, the otoliths (sagittae) of Cook-
eolus also change with adulthood (Fig. 8c, d).
Those of pelagic juveniles are almost perfectly
oval in outline with a ratio of height into length
of about 1:1.25. In adults, the otolith becomes
more elongate (height into length ratio 1:1.6) and
the posteroventral border becomes slightly con-
cave as the marginal ornamentation (lobules)
spreads and the notches deepen. We do not know
of any other perciform in which such differences
in otolith morphology occur with age.

Priacanthus alalaua Jordan and Evermann, 1 904

(Figure 7)

DIAGNOSTIC CHARACTERS. — Second dorsal,
anal and caudal fin membranes red; pelvic fin
membranes jet black; gas bladder with anteriorly
projecting "ears" that extend beyond the body
cavity to the otic bullae, and posteriorly pro-

jecting horns that reach to above the end of the
anal fin (Fig. 9); otoliths (sagittae) with a ventral
keellike blade and centrally positioned pronglike
rostrum (Fig. 8e); preopercle completely scaled;
ctenii filamentous, in rows (Fig. 3b).

MERISTIC DATA.-D. X,13; A. 111,14; P. 18-
19; GR 4-6 + 14-17 = 19-23; pored lateral line
scales 61-66; vertebrae 10 + 13 = 23. Only one
of the 20 specimens we examined for meristic
data had 1 9 elements in the pectoral fin; the rest
had 18.

MAXIMUM SIZE.— The longest individual we
observed was a female 26 1 mm SL (335 mm TL)
from San Benedicto Island, Revillagigedos. The
heaviest individual was a 257 mm SL female
from Socorro Island, Revillagigedos, which
weighed 574 g. Gosline and Brock (1960) report
that in Hawaiian waters P. alalaua "reaches 14
inches in length" (357 mm), but they apparently
did not examine any specimens of this species.

RANGE.— Hawaiian Islands and eastern north
Pacific. In the eastern North Pacific, P. alalaua
has been taken at Alijos Rocks and all of the
Revillagigedo Islands (San Benedicto, Socorro,
Roca Partida and Clarion). In the Revillagigedos,
where sportfi shermen catch fair numbers on oc-

FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS

309

FIGURE 8. Otoliths (sagittae) of eastern Pacific Priacanthidae (otolith length and fish SL, in mm, in parens): a) Pseudo-
priacanthus serrula (7.2 and 2.9); b) Heteropriacanthus cruentatus (5.3 and 195); c and d) Cookeolus hoops (7.6 and 257; 5.5
and 180); e) Pnacanthus alalaua (3.6 and 221). Photos a through d by Jack W. Schott; SEM photo e by Brian White.

casion, it has been taken mostly at night in depths
of "5 to 25 fm or more" (9-46 m).

REMARKS.— Although three individuals of P.
alalaua were taken in gill nets set overnight just
upcoast from Braithwaite Bay, Socorro Island,
in April 1955, they were misidentified as P.
cruentatus, and meristic data from them (e.g.,

"A. 111,13") were reported by Fitch and Schultz
(1978) along with data from seven cruentatus
taken at the same time and place. Fortunately,
their unique otoliths had been removed and
saved, and counts and measurements were made
before they were skeletonized, so subsequent
identification as P. alalaua was easily verified.

310

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19

FIGURE 9. Gas bladder (124 mm total length) from Pria-
canthus alalaua 220 mm SL. Drawing by Mary Butler.

We examined the entire collection (worldwide)
of priacanthids at Los Angeles County Museum
of Natural History and found that arenatus,
hamrur, macr acanthus, meeki, and tayenus pos-
sessed similar peculiar gas bladders and otoliths.
Therefore, we believe these species should be
assigned to Priacanthus.

Heteropriacanthus new genus

TYPE-SPECIES.— Priacanthus cruentatus (Lacepede, 1801).

DIAGNOSIS.— Second dorsal, anal and caudal
fins with rust-colored spotting (dusky in forma-
lin-preserved specimens); pelvic fin membranes
lightly pigmented to clear; gas bladder thin-
walled, contained entirely within the body cav-
ity; otoliths (sagittae) oval in outline with normal
anteroventral rostrum (Fig. 8b); ctenii filamen-
tous, sparse, and only on scale margin (Fig. 3c);
posterior margin of preopercle scaleless.

ETYMOLOGY.— From Greek heteros (different),
alluding to its being different from other Pria-
canthus.

Heteropriacanthus cruentatus (Lacepede, 1801)

(Figure 10)

DIAGNOSTIC CHARACTERS.— As for the genus.

MERISTIC DATA.-D. X.12-13; A. 111,14-15;
P. 17-19; GR 5-6 + 17-20 = 22-25; pored lat-
eral line scales 57-65; vertebrae 10 + 13 = 23.
Only two of the 6 1 specimens we examined for
meristic data had 1 2 dorsal soft rays; the rest had
13. One specimen had an anal count of V,13
(LACM 32283), obviously a freak, so it was not
included. Of the remaining 60 specimens, two
had counts of 111,15 and the rest 111,14. All but
eight specimens had 18 total elements in their
pectoral fins; three had 17 and five had 19.

MAXIMUM SIZE.— The largest individual we
observed (sex undetermined) from the eastern
Pacific was 247 mm SL (3 1 5 mm TL). This fish
(SIO 70- 1 36) was from the Gulf of Chiriqui, Pan-
ama. We did not obtain a weight for it or for any
other large H. cruentatus.

RANGE.— Worldwide in tropical and subtrop-

ical seas. In the eastern Pacific, H. cruentatus
ranges from Guadalupe Island, Baja California
(SIO 60-18) to the Galapagos Islands, Ecuador
(numerous West Coast collections) in depths of
"2 to 15 fm" (3.6-27 m) at least. Within this
range, it has been collected at all of the Revil-
lagigedo Islands, Hurricane Bank, the Tres Mari-
as Islands (Mexico), Cocos Island (Costa Rica),
and many islands off Panama. Its mainland dis-
tribution is not so extensive, ranging from Cape
San Lucas, Baja California (UCLA-W52-259) to
Panama (SIO 70- 140).

REMARKS.— Typically an overall reddish or
crimson when alive, H. cruentatus often will de-
velop silvery marbling or blotching on the sides
and back. Thomson et al. ( 1 979) suggest that this
color pattern results from stress.

Although we examined specimens of//, cruen-
tatus from several localities far removed from
the eastern Pacific, we were unable to find any
differences that could be considered of specific,
or even subspecific, magnitude.

FOSSIL RECORD

Fossil priacanthids have been reported only
from Europe and only from Eocene deposits.
These fossil remains have consisted of skeletal
bones and impressions from Italy (Pristigenys
substriata: see Fritzsche and Johnson 1981), oto-
liths from Belgium and France (Pristigenys rutoti
and P. caduca: Stinton and Nolf 1970; Nolf 1 973),
and otoliths and dorsal fin spines from England
(P. bella, P. spectabilis, and P. dentifer. Stinton
1980).

In identifying fossil fishes, an ideal situation
would be to have a three-dimensional specimen
with all bony elements and conventional char-
acters present (e.g., viscera, gas bladder, scales,
otoliths, etc.). Unfortunately, this does not hap-
pen. Soft parts can only be inferred. Rarely are
three-dimensional fossil fishes found, and even
two-dimensional skeletal impressions are not all
that common compared with isolated teeth,
scales, otoliths, and bones (Schafer 1972). Ob-
viously, the more complete the fossil specimen
or specimens, the greater the likelihood of mak-
ing a correct identification.

The skeletal impressions of P. substriata from
Italy generally have been in excellent condition
and have permitted direct comparison of many
salient features with the same features on extant
priacanthids. None of these two-dimensional
skeletal impressions has contained otoliths, how-

FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS

311

FIGURE 10. Heteropriacanthus cruentatus. Photo by Paul Gregory.

ever; to associate isolated otoliths with the genus
Pristigenys is pure speculation.

In using otoliths (sagittae) as a taxonomic char-
acter, the most important feature for determining
family and/or generic relationships (including
ancestry) is the sulcus or groove on the inner face:
its position, configuration, proportions, angle of
curvature, and other features. If features of the
sulcus do not match exactly, the otolith cannot
belong to the same genus as the one to which it
is being compared. Overall otolith shape, ratio
of height into length, marginal ornamentation
and similar surface features are important pri-
marily at species level.

FOSSIL ACCOUNTS
Pristigenys substriata (Blainville, 1818)

(Figure 1 1)

White (1936) presents an excellent account of
the taxonomy of this species as follows:

Among the numerous fossil fishes described by Volta (1796
. . .) from the lower Lutetian of 'Monte Bolca' was a small
imperfect specimen which he identified with the living species
Chaetodon striatus, . . . illustrating his description with a figure
that is unusually obscure; this specimen was later referred by
de Blainville (1 8 1 8 . . .) to a new species C. substriatus. Agassiz
(1835 . . .) recognizing that the fossil was not a Chaetodon,

renamed it completely, Pristigenys macrophthalmus, and
sketched a few of its more obvious characters, which led him
to suppose that this was 'un genre voisin de Beryx'; and under
Agassiz' name it was listed among the Berycidae by A. S.
Woodward (1901 . . .); finally, Eastman (1905 ...) added de-
tails omitted by previous authors, and re-figured the unique
original specimen. Eastman rightly named the species Pristi-
genys substriatus (for while Agassiz' genus is good, so is de
Blainville's species), and retained it in the Berycidae.

White went on to report that the fossil was not
a berycoid but a priacanthid, which, in his opin-
ion, was identical with the extant Pseudopria-
canthus Bleeker, 1 869, and that this generic name,
by reason of its later publication date, must be
replaced by Pristigenys.

White's report appears to have been over-
looked by subsequent authors until Myers (1958)
called it to the attention of contemporary ich-
thyologists. Subsequently, as discussed by
Fritzsche and Johnson (1981), there has been
considerable controversy as to the validity of
placing Pseudopriacanthus in the synonymy of
Pristigenys. In concurring with such synonymy,
Fritzsche and Johnson point out that a "predor-
sal bone" is a character shared only by Pristi-
genys and Pseudopriacanthus among all priacan-
thid genera. Interestingly, they support their
contention of close relationship by presenting four

312

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19

FIGURE 1 1 .
History).

Pristigenys substriata (BMNH P. 1 6370) 155 mm SL. Photo courtesy of Colin Patterson, British Museum (Natural

sets of morphometric data and two sets of me-
ristic characters (from Pristigenys substriata and
Pseudopriacanthus altus), while one paragraph
later they state that "morphometric characters
are not generally considered to be valid indica-
tors of relationship" within the perciforms.

After carefully examining all available char-
acters for the fossil Pristigenys substriata and for
three species of the extant Pseudopriacanthus (i.e.,
alt us, niphonius, and serrula), it is our contention
that both Pristigenys and Pseudopriacanthus are
valid genera. While we agree that these two gen-
era share at least one character that appears to
be of generic magnitude (i.e., the "predorsal bone"
of Fritzsche and Johnson), there are other salient
characters of equal magnitude that are shared
with other priacanthid genera or are distinctive
within their own genus.

Based upon the associated Monte Bolca fish
fauna, Pristigenys substriata was living in a pe-
lagic environment. The pelagic environment and

extremely long pelvic fins (Fig. 5) are shared with
juvenile Cookeolus, and apparently all known
specimens of P. substriata represent juveniles as
none exceeds 155 mm SL. The scales of Pseu-
dopriacanthus (Fig. 3d) are unique among pria-
canthids for the shape and arrangement of ctenii;
scales of Pristigenys appear to resemble those of
Priacanthus or Heteropriacanthus (Colin Patter-
son, pers. comm.). Other features of Pristigenys
(e.g., scale size, body shape, dorsal fin spine, and
ray lengths) are intermediate to the same features
as found on Cookeolus and Pseudopriacanthus.
There is other less salient evidence to support
retention of both Pristigenys and Pseudopria-
canthus as valid genera.

Pristigenys rutoti (Leriche, 1905)
(Figure 12b)

Otoliths of this species were described and fig-
ured from Belgium by Leriche as "Sparidarum
rutoti." Subsequently, Schubert (1916) reported

FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS

313

FIGURE 12. Otoliths (sagittae) of: a) Pseudopriacanthus ni-
phonius (type species of Pseudopriacanthus) 6.5 mm long; b)

their occurrence in England (refuted by Stinton
1980) and mentioned their similarity to "Oto-
lithus (Sparidarum) rutoti Leriche [sic]," but then
erroneously referred them to Koken's (1891)
Otolithus (Sparidarum) gregarius using the com-
bination "Otolithus (PagellusT) gregarius." Ad-
ditional errors in their taxonomy appeared in
later years by other authors, climaxed by their
being placed in "family Pseudopriacanthidae"
(genus Pseudopriacanthus) by Stinton and Nolf
(1970). Because Leriche used the genitive plural
in giving these a generic name, he cannot be
considered the authority for the species, nor can
the 1905 publication date be considered valid.
Schubert might possibly qualify as the author,
but depending upon interpretation of the Inter-
national Rules of Zoological Nomenclature, this
also is questionable. It is entirely possible that
Stinton and Nolf are the authorities for "Pristi-
genys rutoti,'" with a publication date of 1970,
but the matter is irrelevant to priacanthid tax-
onomy because features of the sulcus of this fossil
otolith do not permit its placement in family
Priacanthidae. In sagittae of extant priacanthids
(except Priacanthus) the upper and lower rims
of the ostium (anterior part of the sulcus) con-
verge posteriorly to appear slightly ovoid and
funnel-shaped (Fig. 7a-d).

In the fossil otolith, the dorsal rim of the sulcus
sweeps posteriorly in a continuous, gentle sig-
moid curve, making it difficult to distinguish os-
tium from cauda without reference to the ventral
rim, which sweeps abruptly dorsad at its juncture
with the cauda. The ostium comprises less than
37% of total otolith length in three species of
Pseudopriacanthus, whereas in Pristigenys rutoti,
ostium length exceeds 41% of otolith length. Fi-
nally, when priacanthid otoliths (except Pria-
canthus) are placed with the outer face down on
a flat surface, the sulcal side is highly arched
(convex). Sagittae of P. rutoti lie almost flat when
placed in this position.

Pristigenys caduca Nolf, 1973
(Figure 12c)

Features of the sulcus of this otolith do not
permit a placement in the family Priacanthidae.

Pristigenys rutoti 2.7 mm long; and c) Pristigenys caduca 2.7
mm long. Photo of Pseudopriancanthus by Jack W. Schott;
SEM photos of Pristigenys by Brian White. Sulcus of fossils
highlighted with broken inked line to show configuration.

314

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 19

Again, the dorsal rim of the sulcus forms a con-
tinuous, gentle sigmoid curve, while the lower
rim of the ostium sweeps abruptly upward at its
posterior terminus— a beryciformlike ostial
character, but also found in such unrelated fishes
as Lactarius, Epigonus, Ambassis, Glaucosoma,
some pempherids, and others.

Pristigenys bella Stinton, 1980

We have not handled otoliths from this species,
but from Stinton's excellent photographs, it ap-
pears that the holotype and paratype shown in
his text figure 34 are valid priacanthid sagittae.
We cannot vouch for their generic placement,
however, as otoliths do not accompany the skel-
etal impressions of 'Pristigenys substriata, the type
of the genus. Again, because of ostial configu-
ration, we question the assignment of the otoliths
(presumably P. bella) shown in Stinton's Plate
1 3 (figs. 27 and 28) to family Priacanthidae. Stin-
ton states that these otoliths "are from immature
fish and demonstrate the ontogenetic changes
[found in otoliths of these fishes]." Except for
Cookeolus, which goes from a pelagic existence
as juveniles to a benthic existence as adults, we
have not observed ontogenetic changes in pria-
canthid sagittae, and the changes in Cookeolus
otoliths are not the same as those attributed by
Stinton to his fossil P. bella.

Pristigenys spectabilis Stinton, 1980

We have examined a dozen otoliths from this
species, and judged by sulcal characters, it should
not be assigned to family Priacanthidae. Stin-
ton's excellent photographs of type material also
show the non-priacanthid ostial configuration that
precludes their being priacanthids.

Pristigenys dentifer Stinton, 1980

Based upon Stinton's photographs of the oto-
liths of P. dentifer, we believe that more than
one species is involved. Some of his illustrated
sagittae are very similar to priacanthid otoliths
in sulcal characters, while others do not appear
to be. One of his paratypes was found associated
with "a few bones and spines" inside the cavity
of a gastropod, Clavilithes macrospira. Accord-
ing to Stinton "some [of the] peculiarly prickly
fin spines . . . were independently identified as a
species of Pristigenys."

We were loaned one of these fin spines by Colin
Patterson (BMNH), who informed us (pers.

comm.) that these prickly spines "are certainly
different from those of Pristigenys substriata,
which are smooth apart from longitudinal ribs."
We compared the fossil spine with fin spines of
all extant genera of priacanthids and could find
no agreement with any of them. Camm Swift
(LACM) examined the spine and suggested it
bore resemblance to some beryciform fin spines,
but was unable to suggest a family or generic
affiliation.

Otoliths (especially sagittae) generally are ex-
cellent taxonomic tools, but when working with
fossils, factors other than those observed on the
otoliths themselves must be considered. Zooge-
ography, environment and habitat preferences
are especially important. If zoogeography had
been considered, it is doubtful that embiotocid
perch (presently restricted entirely to the North
Pacific Ocean) and Leuresthes, Atherinops, and
other New World atherinids would have been
reported from the tropical and subtropical Eocene
of Europe.

Except for the early pelagic stage of Cookeolus
boops, all extant priacanthids inhabit areas of
high relief. Fishes living in these kinds of habitats
frequently fossilize, but their remains rarely con-
tain otoliths. Fossil deposits that contain otoliths
almost invariably represent faunas that inhabit
flat relief, or pelagic and mesopelagic realism.
The occasional otolith from an inhabitant of
rocky, high-relief habitat found in a fossil deposit
generally represents a prey item or a straggler
into the flat-relief area, a not uncommon phe-
nomenon today. Such otoliths are rare, however,
so the abundance of Eocene sagittae assigned to
family Priacanthidae (Pristigenys spp.) by Eu-
ropean paleontologists indicates that habitat
preference was not a consideration. As already
pointed out, features of the sulcus also were over-
looked when making such assignments. Regard-
less, otoliths are excellent taxonomic tools, ex-
tremely abundant in the fossil record, and if
properly used, can furnish an insight into the past
that can not be gained any other way.

ACKNOWLEDGMENTS

As with any such project, our investigation
could not have been completed without our bor-
rowing or making use of specimens, library ma-
terials, work space, ideas, and the special talents
of others. Our sincere thanks are extended to
Lillian J. Dempster and W. I. Follett (CAS); Paul

FITCH & CROOKE: REVISION OF EASTERN PACIFIC CATALUFAS

315

A. Gregory, Jack W. Schott and Donald L.
Schultze (DFG); Mary Butler, Robert J. Laven-
berg, Camm C. Swift and Brian N. White
(LACM); Richard H. Rosenblatt (SIO); Boyd W.
Walker (UCLA); William A. Bussing (Univ. Cos-
ta Rica); John E. Randall (Bishop Museum, Ha-
waii); Wayne J. Baldwin (Hawaii Institute of Ma-
rine Biology); Bruce B. Collette (National Marine
Fisheries Service, Washington, D.C.); Frederick
H. Berry (NMFS, Miami); Richard W. Hud-
dleston (Chevron Oil Field Research Co.); James

B. Shaklee (now CSIRO, Australia); Peter Forey
and Colin Patterson (British Museum Natural
History); Hans-J. Paepke (East Germany); Chris-
tine Karrer (West Germany); Dirk Nolf (Bel-
gium); F. C. Stinton (England); George Coates
(Australia); and Lee Stockland (Tustin, Calif.).

Wayne C. Starnes (USNM) and Ronald A.
Fritzsche (Humboldt State Univ.), who also are
conducting research on priacanthids, were ex-
tremely helpful.

Finally we wish to thank the skippers and crews
of the Qualifier 105, Red Rooster and Royal Po-
laris and in particular George Cargal and Nor-
man Kagawa.

LITERATURE CITED

AGASSIZ, J. L. R. 1835. Revue critique des poissons fossiles

figures dans 1'Ittiolitologia Veronese. Neues Jahrb. Miner.,

p. 290-316. (Not seen, ref. in White 1936.)
ANDERSON, W. D., JR., D. K. CALDWELL, J. F. McKiNNEY, AND

C. H. FARMER. 1972. Morphological and ecological data

on the priacanthid fish Cookeolus hoops in the western north

Atlantic. Copeia 1972(4):884-885.
BLAINVILLE, H. M. D. 1818. Poissons fossiles (In Nouveau

Dictionnaire d'Histoire Naturelle . . . Nouvelle edition, vol.

27, Paris). (Not seen, ref. in White 1936.)
BLEEKER, P. 1869. Neuvieme notice sur la faune ichthyolo-

gique du Japon. Versl. Akad. Amsterdam (2), 3:237-259.
BLOCK, M. E. 1 792. Naturgeschichte der auslandischen Fische.

Berlin Vol. 6, iv + 126 p.
EASTMAN, R. 1 905 . Les types des Poissons fossiles du Monte

Bolca au Mus6um d'Histoire naturelle de Paris. M6m. Soc.

geol. France 13:1-31.

FITCH, J. E., AND R. J. LAVENBERG. 1975. Tidepool and near-
shore fishes of California. Univ. Calif. Press, Berkeley. 156 p.
FITCH, J. E., AND S. A. SCHULTZ. 1978. Some rare and unusual

occurrences of fishes off California and Baja California. Calif.

Fish Game 64(2):74-92.
FORSSKAL, P. 1775. Descriptiones animalium avium, am-

phibiorum, piscium, insectorum, vermium; quae in itinere

oriental observavit. Post mortem auctoris edidt Carsten Nie-

buhr. Copenhagen. 20 + XXXIV + 164 p.
FRITZSCHE, R. A. 1978. The first eastern Pacific records of

bulleye, Cookeolus hoops (Bloch and Schneider, 1 80 1 ), (Pisces,

Priacanthidae). Calif. Fish Game 64(3):2 19-221.
FRITZSCHE, R. A., AND G. D. JOHNSON. 1981. Pseudopria-

canthus Bleeker, a synonym of the priacanthid genus Pris-

tigenys Agassiz. Copeia 1981(2):490-492.
GILBERT, C. H. 1891. A supplementary list of fishes collected

at the Galapagos Islands and Panama, with description of

one new genus and three new species. U.S. Natl. Mus., Proc.

13:449-455.
GOSLINE, W. A., AND V. E. BROCK. 1960. Handbook of

Hawaiian fishes. Univ. Hawaii Press, Honolulu, ix + 372 p.
JORDAN, D. S., AND B. W. EVERMANN. 1904. Descriptions of

new genera and species of fishes from the Hawaiian Islands.

U.S. Fish. Comm., Bull. 22:161-208.
KOKEN, E. 1891. Neue Untersuchungen an tertiaren Fisch-

Otolithen II. Deutsch. Geol. Ges., Zeitsch. 43:77-170.
LACEPEDE. 1 80 1 . Histoire naturelle des poissons, Paris. Vol.

3, 558 p.
LERICHE, M. 1 905. Les Poissons eocenes de la Belgique. Mu-

see Royal Hist. Nat. Belg., Mem. 3:49-228.
MYERS, G. S. 1958. The priacanthid fish genus Pristigenys.

Stanford Icthyol. Bull. 7:40-42.
NOLF, D. 1973. Deuxifcme note sur les te!6osteens des sables

de Lede (Eocene Beige). Bull. Soc. beige Geol., Paleont.,

Hydrol. 81(l-2):95-109.
OKEN, L. 1817. Cuviers und Okens Zoologien neben einander

gestellt. Isis oder Encyclopaedische Zeitung 8(148):1181-

1183.
SCHAFER, W. 1972. Ecology and palaeoecology of marine

environments. Univ. Chicago Press, xiii + 568 p.
SCHNEIDER, J. G. 1 80 1 . M. E. Blochii. Systema ichthyologiae

. . . post obitum auctoris correxit, interpolavit J. G. Schnei-
der. Berlin, Ix + 584 p.
SCHUBERT, R. J. 1916. Obereocane Otolithen vom Barton

Cliff bei Christchurch (Hampshire). Kaiserlich-kSniglichen

Geologischen Reichsanstalt, Jahrb. 65(3-4):277-289.
STINTON, F. C. 1980. Fish otoliths from the English Eocene.

Part IV. Palacontographical Soc. Monogr., p. 191-258.
STINTON, F. C., AND D. NOLF. 1970. A teleost otolith fauna

from the Sands of Lede, Belgium. Bull. Soc. beige Geol.,

Paleont., Hydrol. 78(3-4):2 19-234.
THOMSON, D. A., L. T. FINDLEY, AND A. N. KERSTITCH. 1979.

Reef fishes of the Sea of Cortez. John Wiley & Sons, New

York. XV + 302 p.
VOLTA, G. S. 1796. Ittiolitologia Veronese del museo Boz-

ziano ora annesso ... 2 vols. Verona. (Not seen, ref. in White

1936.)
WHITE, E. I. 1936. On certain Eocene percoid fishes. Ann.

Mag. Nat. Hist., ser 10, 18(103):43-54.
WOODWARD, A. S. 1901. Catalogue of fossil fishes in the

British Museum Natural History. Part IV, London. 636 p.

(Not seen, ref. in White 1936.)

CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, CA 941 18

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Vol. 43, No. 20, pp. 317-321, 3 figs.

December 11, 1984

AMAZONSPRATTUS SCINTILLA, NEW GENUS AND SPECIES

FROM THE RIO NEGRO, BRAZIL, THE SMALLEST

KNOWN CLUPEOMORPH FISH

By
Tyson R. Roberts

California Academy of Sciences, Golden Gate Park, San Francisco, California 94118

ABSTRACT: Amazonsprattus scintilla new genus and species, inhabiting the Rio Negro and its tributaries in
the Amazon basin of Brazil and feeding on minute aquatic Diptera and planktonic Cladocera, is the smallest
known clupeomorph fish. The largest specimen is 19.5 mm standard length, but males and females are
sexually ripe as small as 14-16 mm. Superficially it looks very much like a herring and particularly resembles
the tropical western Atlantic genus Jenkinsia (Clupeidae). On the other hand, further investigation might
indicate that it is closely related to some small and poorly known Amazonian anchovies currently placed in
Anchoviella (Engraulididae).

INTRODUCTION

Among the secondary freshwater fishes sharing
the Amazon basin with 1500 or so species of
primary freshwater Ostariophysi are somewhat
more than a dozen species of the clupeomorph
families Clupeidae and Engraulididae. Amazo-
nian Clupeidae, or herrings, include several
species of Ilisha or Pellona, some piscivorous
and attaining nearly one meter; Pristigaster cay-
ana Cuvier, 1 829, with its greatly expanded tho-
rax, attaining perhaps 250 mm; and Rhinosar-
dinia amazonica (Steindachner, 1880), typically
40-60 mm long. The Engraulididae, or ancho-
vies, include Cetengraulis juruensis Boulenger,
1898, attaining perhaps 200 mm, and a number
of species currently placed in Anchoviella, some
as small as 30-40 mm. Recently, while searching
for comparative material of larvae, I was thus
surprised to find some sexually ripe Amazonian
fishes, which looked like clupeomorphs, less than

20 mm in standard length. Study of cleared and
stained preparations confirmed that these spec-
imens are not Ostariophysi and that they rep-
resent an undescribed genus and species of
clupeomorph.

Amazonsprattus, new genus

TYPE-SPECIES.— Amazonsprattus scintilla, new species.

DIAGNOSIS.— Minute, slender, scaleless clu-
peomorphs without pre- or post-pelvic abdom-
inal scutes. Ventral myotomic progression in-
complete. Premaxilla absent or minute and
toothless. Maxilla with 1 6-20 very small conical
teeth. Two supramaxillae. Dentary and palate
usually toothless (one tooth observed on dentary
in one specimen). Branchiostegal rays 4-5. Dor-
sal fin with 12-13 rays, its origin in posterior half
of body; anal fin with 14-16 rays, its origin below
anterior third of dorsal fin. Pectoral fin rays 7-
9. Pelvic fin rays 6.

[317]

318

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 20

FIGURE 1. Amazonsprattus scintilla, 17.0 mm (holotype, CAS 52175).

Amazonsprattus scintilla, new species

(Figures 1-3)

HOLOTYPE.— CAS 52175, 17.0 mm (sex undetermined), Rio
Jufari between Castanheiro Grande and Santa F6, collected by
Martin Brittan, 21 April 1964.

PARATYPES.— CAS 52176, 18: 14.3-19.0 mm (five males,
eight females, six sex undetermined), collected with holotype
(5: 16.5-1 9.0 mm cleared and stained with alcian and alizarin);
CAS-SU 68891, 19.5 mm (sex undetermined), Rio Negro at
Santa Isabel, collected by Carl Ternetz, 17 January 1925.

DESCRIPTION.— Head compressed and mod-
erately elongate, its length almost four times in
standard length. Eyes moderately large and
strongly compressed or flattened laterally. Eye
diameter about four times in head length. Entire
medial surface of eyes closely approximate (so
that forebrain is confined to a small space dorsal
to eyes), and medial surface of eyes just as flat
as lateral surface. Ventral surface of eyeball with
a prominent choroid fissure. Hyaline eyelid well
developed. Snout moderately elongate, its length
about equal to eye diameter. Nasal organ mod-
erately large, with rather small anterior and pos-
terior nostrils. Mouth terminal. Lower jaw elon-

gate, extending posteriorly to below posterior
margin of eye, but gape much restricted and en-
tirely anterior to eye. Maxilla slender and mod-
erately elongate, extending posteriorly to below
anterior margin of eye. Anterior and posterior
portions of cranial fontanel open, posterior por-
tion entirely bridged by a narrow, cartilaginous
taenia medialis. Circumorbital bones apparently
five (very fragile, hence easily broken, and stain-
ing very weakly with alcian). Hyosymplectic en-
tirely cartilaginous, without large, axe-shaped
anterodorsal process usually present in clupeids.
Opercle with strongly concave dorsoanterior and
dorsoposterior margins; dilatator process of
opercle present but weakly developed. First gill
arch with a total of 18-19 large, elongate gill
rakers on its leading edge. Rakers on trailing edge
of first gill arch and on successive arches slightly
less numerous and considerably smaller. Upper
and lower pharyngeal toothplates small, with 8-
10 conical teeth (branchial and hyoid arches oth-
erwise edentulous). Hyoid and branchial arches
entirely cartilaginous except for branchiostegal
rays, gill rakers, and pharyngeal toothplates.

hyosymplectic

palatine

maxilla

supramaxillae

opercle

lower jaw

preopercle
Meckel's cartilage

subopercle
interopercle

1 mm

FIGURE 2. Amazonsprattus scintilla, 18.0 mm, CAS 52176. Lateral view of jaws, suspensorium, and opercular bones (hy-
osymplectic and palatine cartilaginous).

ROBERTS: MINUTE AMAZONIAN CLUPEOMORPH

319

FIGURE 3. Amazonsprattus scintilla, 18.8 mm, CAS 52176. Axial skeleton. Total vertebrae 17 + 21 = 38; pectoral girdle
abnormally incomplete; a series of 10-1 1 small, irregularly triradiate intermuscular bones lateral to vertebrae 2-13 has been
omitted for clarity.

Body compressed, slender, its greatest depth
about 6-8 in standard length. Abdomen round-
ed. Myotomes well denned, those fully formed
33-34, or about four fewer than total vertebrae.
Dorsal and anal fin rays low set, with moderately
falcate margins, dorsal fin rays about 1.5 times
longer than anal. Pectoral and pelvic fins rela-
tively small, pelvic smaller than pectoral. Pelvic
scute variably developed, absent or failing to stain
in some specimens, weakly to strongly stained
with alcian or alizarin in others. In specimens
with relatively well developed scutes there is an
elongate anteromedian process and an elongate
lateral ascending process on each side. Pectoral
girdle with bony posttemporal, supracleithrum,
and cleithrum (postcleithra absent), cartilaginous
scapulocoracoid, and three rows of radials (prox-
imal, medial, and distal, with five, five, and eight
radials respectively). Caudal fin moderately
deeply forked, upper and lower lobes about equal
and with rounded margins. Upper and lower lobes
overlapping slightly when adducted. Principal
rays 4-5 of upper lobe and 3-4 of lower lobe
with delicate alar flaps (not illustrated). In an
18. 8 -mm specimen the large alar flap on ray 5
consisted of four overlapping scalelike laminae
(lightly stained with alcian) that may actually be
modified scales. Caudal fin with 10 + 9 principal
rays, 8-9 upper and 8 lower procurrent rays. Cau-
dal fin skeleton with a parhypural, six separate
hypurals, and a single epural. Hypural 2 fused to
complex ural centrum (as in many clupeoids).

Total vertebrae 37(4) or 38(1), 16-17 abdom-
inal plus 20-2 1 caudal. All vertebrae with a sim-
ple, slender neural spine. Supraneurals 6-8. Or-
igin of pelvic fin below vertebrae 13-14, of dorsal
fin above vertebra 18, and of anal fin below ver-
tebrae 20-21. All abdominal vertebrae except
first two with fully developed ribs. First two ver-

tebrae usually without ribs, sometimes second
vertebrae with incompletely developed ribs (Fig.
3). Distal ends of most abdominal ribs deflected
posteriorly. First two caudal vertebrae with re-
duced ribs. Intermuscular bones well developed
anteriorly and posteriorly. Anteriorly two dis-
similar and morphologically complex sets of in-
termuscular bones. A series of about 13 epipleu-
ral intermuscular bones parallel to abdominal
vertebrae 3-16. Anteriormost eight epipleurals
with a well developed anteromedial process
proximally; this process is absent from last five
epipleurals, which become progressively smaller.
Distal ends of epipleurals, except reduced pos-
teriormost one, closely approximated to distal
half of ribs (Fig. 3). In addition to epipleurals, a
series of 10-11 small, irregularly triradiate in-
termuscular bones lies just dorsal to epipleurals
and directly lateral to centra of abdominal ver-
tebrae 2-12 or 13 (not illustrated). Posterior two
processes of these triradiate elements lie quite
near body surface, but anterior process lies much
deeper. Posteriorly, two similar series of 8-10
simple dorsal and ventral intermuscular bones
extend laterally just above and below caudal ver-
tebrae 9-19 (Fig. 3).

Alimentary canal with a well-defined stomach.
Pyloric caeca in two or three groups: a dorsal
group with one or two elongate caeca; a ventral
group with about four or five elongate caeca; and
sometimes a second ventral group of about four
short, poorly defined or only partially separate
caeca. Intestine straight. Gut contents of four
specimens were as follows: specimen 1) two dip-
teran pupae; specimen 2) a single dipteran pupa;
specimen 3) numerous small Cladocera of two
size classes, 270 x 1 50 ^m and 72 x 55Mm;and
specimen 4) a single dipteran pupa, several small
dipteran larvae, and moderately numerous

320

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43, No. 20

cladocerans of a single kind. My sketches of the
cladocerans in this fourth specimen, 424 x 255
^m, with a pair of curved, divergent, strongly
deflected horns 1 84 nm long projecting from the
rostrum, were tentatively identified as Bosmi-
niopsis deitersi Richard, 1 895, by Thomas Zaret.

Gonads readily identifiable in most of the type-
specimens. Eight, 15.9-18.2 mm, have creamy
or pale orangish ovaries with eggs in more or less
good condition observable through body wall with
transmitted light. One of these, 17.3 mm, con-
tained 20 eggs 0.2 mm in diameter. Five, 14.3-
16.2 mm, have milk-white testes. In two of these
the testes are particularly well developed and ex-
hibit numerous "segments" or laminae, about
five per myotome, comparable to laminae ob-
served in testes of other minute teleosts (e.g.,
Sundasalarvc, Roberts 1 98 1 , fig. la). In both sexes
the gonads appear to be single and occupy only
the posterior half of the body cavity, from about
the origin of the pelvic fin to the vent. In seven
specimens, 16.0-19.5 mm, gonads not observed
and sex undetermined.

Judging from preserved material live Ama-
zonsprattus probably are translucent or even
transparent. Only a few melanophores on head,
largest and most obvious a group of about six
superficial to cleithrum and clearly visible through
gill cover, and two large ones on either side of
dorsoposterior margin of hindbrain (Fig. 1). A
number of large, deep-lying melanophores as-
sociated with postero ventral portion of cranium
(not illustrated). Dorsal surface of cranium oth-
erwise usually devoid of pigment. Tip of snout
and lower jaw, and side of head just below eye
and midway between eye and end of gill cover
sometimes with a few small melanophores. Body
with relatively few melanophores, mostly on
ventral half, including base of anal fin and caudal
peduncle. A row of about 10 melanophores, one
per segment, along each ventral myotomic bor-
der, and a midventral row of about five near
pelvic fins. Two rows of segmental melanophores
near anal fin base: one row at ventral end of
myotomes and between pterygial muscles,
another on bases of anal-fin rays. A row of small
melanophores, more than one per segment, on
ventral portion of caudal peduncle, and some
small melanophores near base of caudal fin. A
nearly straight row of small segmental melano-
phores on side of body just above midline (with-
out evident anatomical relationship to any un-

derlying structures). In holotype and most
paratypes this row commences posterior to dor-
sal fin origin (Fig. 1), but in some paratypes it
extends nearly entire length of body. Dorsal sur-
face of body devoid of pigmentation. A few me-
lanophores on basal portion of anterior dorsal
fin rays, but dorsal fin without basal melano-
phores like those of anal fin. Caudal fin relatively
densely pigmented, with large melanophores
more or less regularly distributed on upper and
lower lobes (Fig. 1 ); an area near middle of caudal
fin devoid of melanophores. Sexual differences
in pigmentation not observed.

NOTE ON TYPE-LOCALITIES.— The Rio Jufari is
a low-gradient, swampy tributary with an enor-
mous mouth-bay at its confluence with the Rio
Negro, about 20 km upriver from the relatively
narrow mouth of the much more important Rio
Branco. Maps I have seen do not show Castan-
heiro Grande or Santa Fe;*according to the col-
lector (pers. commun. M. Brittan, March 1983)
several days of slow boat travel up the Jufari were
required to reach the collecting sight. Santa Is-
abel is an old name for the modern town of Ta-
purucuara. This portion of the Amazon basin lies
within equatorial rain forest; here the waters of
the Rio Negro and its tributaries (excepting the
Rio Branco) are generally darkly tinted, have a
pH of 4-5, and are extremely low in mineral
conduct.

ETYMOLOGY.— Amazon; and sprattus (Latin,
masc.), a herring or herringlike fish; scintilla (Lat-
in, masc.) a spark, hence the smallest trace or
particle (employed as a noun in apposition).

DISCUSSION

Few collectors have preserved specimens of
the smallest Amazonian fish species, and it may
well be that the smallest species of various other
groups have yet to be discovered. In the Amazon,
where community structure of freshwater fishes
may be more complex than anywhere else, mi-
nute body size seems to have arisen primarily in
response to biotic factors. This, in my opinion,
is the general reason why so many of the smallest
Amazonian fishes belong to secondary freshwa-
ter fish groups which in terms of relative numbers
of species represent an insignificant fraction of
the fauna. Amazonsprattus provides an excellent
example of the survival of a group of secondary
fishes in the midst of the Amazonian ichthy-
ofauna by evolution of minute body size and an

ROBERTS: MINUTE AMAZONIAN CLUPEOMORPH

entirely freshwater life history, possibly involv-
ing year-round continuous reproduction of mi-
nute young. Other particularly striking examples
of the phenomenon include the two minute
species of Poecilia (or Pamphorichthys) and the
two or more minute species of Microphilypnus.
These are the only members of the families Poe-
ciliidae and Eleotrididae inhabiting the interior
of the Amazon basin. For further discussion of
these and other examples see Roberts (1972).

Relationships of Amazonsprattus are unclear,
and for the moment it is perhaps best left un-
assigned to family. Having examined its skeletal
anatomy and made comparisons with a number
of herrings and anchovies, I was inclined to place
the genus in Clupeidae, largely on account of its
jaw structure, which is unlike anything I have
observed in Engraulididae. But my colleague
Gareth Nelson, who is studying Amazonian An-
choviella and has examined some small unde-

321

scribed species I have not seen, is inclined to
believe that it may belong to this group of En-
graulididae.

ACKNOWLEDGMENTS

For information, comments, or other assis-
tance I wish to thank Martin Brittan, Norma
Chirichigno, Dusty Chivers, Lillian Dempster,
William N. Eschmeyer, Karsten Hartel, Michael
Hearae, Albertina Kameya, Vincent Lee, Sarah
Ward, Thomas Zaret, and George Zorzi. The
manuscript was reviewed by Gareth M. Nelson
and Peter J. Whitehead.

LITERATURE CITED

ROBERTS, T. R. 1972. Ecology of fishes in the Amazon and
Congo basins. Bull. Comp. Zool. 1 43(2): 1 17-147.

. 1981. Sundasalangidae, a new family of minute fresh-
water salmoniform fishes from Southeast Asia. Proc. Cali-
fornia Acad. Sci. 42(9):295-302, 6 figs.

CALIFORNIA ACADEMY OF SCIENCES

Golden Gate Park
San Francisco, California 94 1 1 8

INDEX TO VOLUME 43

(Compiled by Lillian J. Dempster)
New names in boldface type

Acanthocybium solanderi 301
Acarina 14
Acestrorhynchus 206
Acridium

coloratum 49-50

xanthopterum 44, 5 1
Ahlia 60

egmontis 64
Albula chinensis 210
lAlbula chinensis 210
Alloeocarpa 239

sp. 239
Allopora 245
Alopias

superciliosus 89

vulpinus 89
Alopiidae 94-96
Alytes obstetricans 217
Amazonsprattus 3 1 7-32 1

scintilla 3 17-321
Ambassis 314
Anchoviellalll, 321
Andricus kollari 132
Anthias macrophthalmus 302

^/7/M5 126

figulus 126

Aplidium californicum 244
Aplochitonidae 203, 215
Arctocephalus doriferus 229-230
Argentinoidei 201
Ascidiacea 239
Atherinops 314
Atractoscion nobilis 232
Aulopodidae 201

Balanus

glandula 244

nuW/w 244-245
Balitora 68
Baltimora recta 49
flflriYius 151, 156-157
Bdellostoma

cirrhatum 264

dombeyi 255

Forsteri 264

heptatrema 250, 264
Benthenchelys 60

cartieri 64
Berycidae 311

269-270, 281-282

270-271
tuberculata 270, 281-282
Blakeae 269, 282
Bosminiopsis deitersi 320
Brycon 142

Calophysoides 275

Calyptrella 275

Carabidae 159

Carchariniformes 93

Carcharodon carcharias 89, 107, 221-238

Carinotetraodon 15

lorentziana 49

parqui 49

strigillatum 49
Cetengraulis juruensis 317
Cetorhinidae 94-96
Cetorhinus 95-96, 109-110

maximus 96, 232
Chaenogobius sp. 181

striatus 3 1 1

substriatus 3 1 1
Characoidei 201
C7z<?/a 144
Chelonodon 2
Chlorophthalmidae 201
Chonerhinos 1-16

africanus 1, 4, 7-10

amabilis 1, 3-7, 11-15

modestus 1-15

naritus 1-2, 5

nefastus 1,3-5,9-11, 13-16

remotus 1, 3-5, 9, 11-15

silus 1,3-5,9, 11-16
Chonerhinus 2

africanus 1

modestus 5

naritus 5
Chromacrini 43
Chromacris 43-58

color ata 44-51, 54

colorata group 45, 47, 51

color ata- miles group 50

icterus 44, 46-48, 50, 54-56

m//es 44-48, 51,55

minuta 43-47, 5 1

nuptialis 44-48, 53

[323]

324

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43

peruviana 44, 46-47, 49-50, 54, 56

psittacus 46-48, 50, 54

psittacus pacificus 43-44, 47, 54-55

psittacus psittacus 44-45, 54-55

speciosa 43-53

trogon 46-50, 54-55

trogon group 45, 47, 50-51, 54

trogon intermedia 43-45, 54

trogon trogon 44-45, 53-54
Chthamalus dalli 244
Cladocera317, 319
Clavilithes macrospira 314
Clidemia 269, 271,274

petiolaris 275

saltuensis 274

tetrapetala 269, 271-274

trichosantha 269, 273-275
Clupeidae218, 317, 321
Cnemidocarpa 247
Coleoptera 159
Colomesus 15
Compositae 49
Cookeolus 302, 304-305, 308, 312, 314

boops 301-309, 312,314
Cynoscion nobilis 232

Cyprinidae 67, 142-143, 148, 151-152, 155-157
Cyprinvs

balitora 67, 74

sucatio 67, 69

Dallia pectoralis 199-200
Dalophis rupelliae 20-2 1
Danio 151
Dasyatidae 283-300
Dasyatis 284-289, 292, 295, 298

americana 285

centroura 285-286, 295, 299

garouaensis 283-284, 286, 289-290, 292-295,
298-300

margarita 283-284, 286-290, 292-295, 298-299

margaritella 283-284, 286-295, 298-299

marmorata 285

pastinaca 285-286

rudis 285

sp. 287, 292

ujo 286

ukpam 283-289, 294-300

violacea 285
Dendrodoa 239, 246-247

abbotti 239-248

carnea 239, 245-247

grossularia 245-247

grossularia-carnea-abborti series 247

(Styelopsis) 239, 246-247

(Styelopsis) abbotti 239-248

uniplicata 246-247
Diplomystes 154-155

Diptera 317
Drimys 277

Eleotrididae 321
Elopomorpha 2 1 8
Elops 194, 217-218

hawaiiensis 200, 2 1 8
Engraulididae317, 321
Enhydra lutris 229, 231
Eperlanus 204

chinensis 204-205
Epiactis prolifera 244
Epigonus 314
Epinephelus spp. 30 1
Eptatretus 249-267

burgeri 249

carlhubbsi 249-267

cirrhatus 249-250, 254-261, 264-265

deani 250

heptatrema 264

hexatrema 249

laurahubbsi 249-267

polytrema 262

springer i 254

jtoitf/7 250, 255, 262

strahani 249-267

undescribed 262
Eremopodes sp. 41
Esomus 148
Eugomphodus 94

tricuspidatus 94
Eurystomella bilabiata 244
Exoglossum 143

niphobles 14

Galaxias 194
Galaxiidae 203, 214-216
Galaxioidea 203
Galaxioidei215
Galeocerdo cuvier 89, 229
Galeomorphii 93
Galeorhinus zyopterus 232
Garra 75
Gastromyzon 75

abingdonii 112-113,115,119
tecfo' 112-113, 115, 119
chathamensis 112-113, 115, 119
chilensis 1 1 9

dbwifii 112-113, 115, 119
elephantopus 112-113
ephippium 1 1 1-1 1 3, 1 1 5-1 20
galapagoensis 113
guntheri 111-113, 115-120
hesterna 119
hoodensis 112-115, 119
macrophyes 1 1 3

INDEX

325

microphyes 112-115, 119

nigra 113

nigrita 111-120

phantastica 112-115, 119

porter i 113-114

sp. 114

vandenburghi 112-115, 119

vicina 112-116, 119

vosmaeri 1 1 1

wallacei 112-114, 119
Ginglymostoma 109
Glaucosoma 314
Glenoglossa 59-66

wassi 59-66
Graffenrieda 269, 275

bella 269, 275-277

galeottii 275

gracilis 275

micrantha 275, 277
Gryllus

locusta miles 5 1

speciosus 47,51
Gymnomuraena marmorata 22
Gymnothorax 18, 20

enigmaticus 17-24

leucacme 17, 22

petelli 17, 20, 22

"petelir form 20-21

reticularis 20-2 1

rueppelliae 17, 19-22

rupelli 20

ruppeli 17, 20

"ruppellf' 21

signifer 17, 21-22

undulatus 22

waialuae 17, 22

Hemisalanginae 206, 210
Hemisalanx 206, 210

prognathus 185, 206, 208, 210
Hemitrygon 287

ukpam 295
Hepsetus 194, 206
Heptatrema cirrata 264
Heptatretus banksii 250, 264
Heterodontiformes 93
Heteropriacanthus 301-315

cruentatus 301, 304-305, 307, 309-31 1
Hexanchus griseus 89
Himantura 284

«#«/«• 290, 292, 294, 298
Homaloptera 75

Hypolophus 284
Hypomesus 194

250, 264
cirrhata 264

7//s/za317
Isistius 110

brasiliensis 110
I sums spp. 233

Jenkinsia 317

Lactarius 314
Lamnidac 94-96
Lamniformes 89, 93
Leguminosae 49
Lepidogalaxias 194-195,214-217

salamandroides 2 1 5-2 1 6
Leptasterias pusilla 244
Leucosoma 206, 210

chinensis 205, 210

reeves/ 206, 210
Leuresthes 314
Lipara lucens 136
Lovettia 215
Ludobrama 152-153
Luciosoma 143
Lycium 49

cestroides 49
Lycodontis 20

Macrochirichthys 144, 151
Macrocystis pyrifera 232
Mallotus2l5
Medicago saliva 49
Megachasma 87-1 10

pelagios 87-110
Megachasmidae 87-1 10
Melastomataceae 269-282
Metasalanx 206

coreanus 206, 210

269, 275, 277, 279

arboricola 269, 277-279, 281

mollicula 279
Miconieae271, 277
Microdonophis fowleri 23
Microphilypnus 321
Mirounga angustirostris 229-230
Mitsukurina 105, 107

owstoni 94

Mitsukurinidae 94, 96
Mixodigma leptaleum 110
Mixodigmatidae 110
Monotreta 2,15

Hylaeus pect oral is 136
Hymenoptera 27, 123

interrupta 17, 22
umbrofasciata 17, 21
xanthopterus 22

326

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43

Muraenichthys 60, 62
Muraenidae 18
Mycteroperca spp. 301
Myctophidae 200
Myliobatis californica 230-232
Myrophinae 59-60
Myrophis 60

frio 65
Myxine 25 1-252, 254

glutinosa 255

Nebria 159-177

acuta acuta 167

acuta sonorae 159, 167

altisierrae 159-161, 169, 171, 176

arkansana 167

arkansana arkansana 167

arkansana edwardsi 167

arkansana fragitis 159, 167

arkansana uinta 159, 167

calva 159, 164, 170-171, 173, 177

campbelli 159, 161, 169, 171, 176

crassicornis 166-167

crassicornis intermedia 159, 166

fragilis 159, 167

fragilis fragilis 167

fragilis teewinot 159, 167

gebleri albimontis 159, 163, 169, 171, 173, 177

haida 159, 162, 169, 176

intermedia 159, 166-167

Jeffrey! 159, 162, 169, 176

labontei 159, 163-164, 170, 177

louiseae 159, 162-163, 169, 176

piute 167-168

piute piute 159, 167

piute sevieri 159, 164-165, 168, 170, 172, 174,
177

piute utahensis 159, 168

sierrablancae 159, 164, 170-171, 174, 177

sonorae 159, 167

steensensis 159, 165, 170, 172, 175, 177

trifaria 167-168

trifaria catenata 164

trifaria coloradensis 165

trifaria pasquineli 159, 165-166, 170, 172, 175,
177

trifaria piute 159, 167-168

trifaria tetonensis 159, 167

trifaria trifaria 159, 164-165, 167

trifaria utahensis 159, 168

utahensis 168

virescens 160-161

wallowae 159, 161-162, 169, 171, 173, 176
Nebriini 159-177
Neenchelys 59-60, 62-64

buitendijki 64-65

daedalus 59-66

microtretus 62, 64-65

parvipectoralis 64

spp. 64

Nemamyxine 251-254
Neomysis sp. 181
Neomyxine 251, 254

Neosalanx\%2, 184, 186, 198,201-205,210-212,214,
219

andersoni 179, 181-182, 184, 186, 209, 211-212,
219

brevirostris 179,181,186-187, 209, 2 1 1-2 1 2, 2 1 9

hubbsi2\2

jordani 179-181, 183-186, 190-192, 195, 197, 199,
201-202,204,209-212,219

regani 2 1 2

reganius 179, 181, 186, 202, 212

tangkahkeii taihuensis 212, 219
Neoscopelidae 200
Neomacheilus 12
Notomyxine 254
Notoplana acticola 244
Novumbra hubbsi 217

Odontaspididae 94, 96
Odontaspis 94, 96
Ophichthidae 23, 59-60
Ophichthinae 60
Ophichthus

erabo 17, 23-24

garretti 23

retifer 17, 23

Ophiodon elongatus 231-232
Opsariichthys 154-155
Orectolobiformes 93
Orthoptera 43

Osmeridae 190, 199-201, 203, 215-216
Osmeroidea 179, 203
Ostariophysi 180-181, 317
Osteochilus 154
Otolithus

(Pagellusl) gregarius 3 1 3

(Sparidarum) gregarius 3 1 3

(Sparidarum) rutoti 313
Oxygaster 144

Pamphorichthys 321
Pangasius 13

polyuranodon 13-14
Paramyxine 251, 253, 255
Paraprotosalanx 204-205, 211

andersoni 205
Parasalanx 202, 206-207, 210

acuticeps 206, 208

angusticeps 207-208, 210

annitae 206-208

ariakensis 210

cantonensis 207-208, 210

gracillimus 206-208, 210

INDEX

327

longianalis 206-208

(Salanx) 202

tParasalanx gracillimus 207
Parexoglossum 143
Pellonal\l
Petrolisthes 244
Petromyzon drrhatus 264
Phoca vitulina 236
Pisces 203, 301
Platycara 68

Plecoglossidae 200, 203, 215
Plecoglossus 203, 215
Plecoptera 14
Plectognathi 14
Podocarpus 277
Poecilia 321
Poeciliidae 321
Potamotrygon 283

garouaensis 292
Potamotrygonidae 283
Priacanthidae 301-315
Priacanthus 301-302, 310, 312-313

alalaua 301-303, 305, 307-310

arenatuslQl, 310

cruentatus 301-302, 304, 309-310

hamrurlQl, 310

macracanthus 301, 310

/w**i301, 310

tayenusWl, 310
Prionace glauca 228
Pristigaster cayana 3 1 7
Pristigenys 30 1 , 304, 3 1 1-3 1 4

fc?//a301,310, 314

caducallQ, 313

dentiferlOl, 310, 314

macrophthalmus 3 1 1

rutoti 3 10, 312-313

spectabilis 310, 314

spp. 314

substriata 301, 304, 310-312, 314

substriatus 3 1 1
Protosalanginae 179, 181, 185, 189-190, 192, 194, 198,

202,204,206,211

Protosalanx 179, 182-185, 189, 192, 194, 198-199,
202, 204-205, 211, 214, 216, 218

andersoni 205, 209, 211

brevirostralis 212

brevirostris 209, 211-212

chinensis 179-181, 183-185, 187, 192-193, 196,
199, 201-202, 205, 208, 210-211, 214

hyalocranius 205, 211-212

tangkahkeii 2 1 1
1 Protosalanx tangkahkeii 212
Prototroctes 203
Prototroctidae 201
Pseudocarcharias 105

kamoharai 94
Pseudocarchariidae 94, 96

Pseudomyrophis 59-60, 64-65

atlanticus 65

micropinna 65

nimius 65

spp. 64-65

Pseudopriacanthidae 3 1 3
Pseudopriacanthus 302, 304, 311-313

serrula 301-305, 307, 309, 312

Psilorhynchidae 67

Psilorhynchus 67-70, 73, 75
balitora 67-76
balitora-like species 68
gracilis 67-76
homaloptera 67-69, 72, 75
homaloptera rowleyi 67
pseudecheneis 67-69, 72
swotf/o 67-70, 72-75
sucatio var. damodarai 73-74
variegatus 68, 74

Quercus spp. 277

/?fl/a pastinaca 286
Rasbora 151
Reganisalanx 206

normani 206-208, 210
1 Reganisalanx brachyrostralis 210
Retropinnidae 201, 203, 215
Rhiniodon 109-110

fy/ms 89, 96
Rhinosardinia amazonica 3 1 7

latipennis 44, 53

m/fes 52

m/fes Var. C 52

nuptialis 53

opulenta 44, 55-56

/>«fcs 44, 49-50

peruviana 56

spedosa 52

stolH44, 51

frogort 53

Romalea psittacus 54
Romaleidae 43
Romaleinae 43, 47
136

Sagraea 274

Salangichthyinae 179, 181-182, 186, 189-190, 192,

194-195, 198, 202,204,211
Salangichthys 184, 186, 202-203, 211-213

ishikawae 179-181, 185-186, 199, 204, 209, 213
kishinouyei 2 1 3

microdon 179, 181-187, 201-202, 204, 209, 212-
215,218

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 43

Salangidae 179, 181-183, 185, 194-195, 198-206,210-

211,213-218
Salanginae 179, 181-182, 185, 189-190, 192-195, 198,

204-206, 210, 218
Salangoidea 179-200
Salanx 179, 184, 201-204, 206-207, 210-211

acuticeps 206

argentea 211

ariakensis 181, 185-186, 206-208, 210, 213

brachyrostralis 210

chinensis 181, 187, 204-205, 210, 216

cuvieri 180-182, 186, 188, 192, 194, 197, 199,
202, 206-208, 210

(Hemisalanx) 186, 210

(Hemisalanx) prognathus 179, 210

hyalocranius 204-205, 208, 212

(Leucosomd) 186, 198, 210

(Leucosomd) reevesi 179, 198

(Leucosomd) reevesii 210

prognathus 181, 185-186, 189, 195, 208, 210

reevesi 179, 181, 186, 208, 210

reevesii 210

(Salanx) 186, 202, 207, 210

(Salanx) ariakensis 179, 206

(Salanx) cuvieri 179, 207
tSalanx

argentea 212

brachyrostralis 210

cuvieri 206
Salmacina 244
Salmo 194

Salmonidae 200-201, 203, 214
Salmoniformes 179, 181, 195, 199-200, 214-215, 218
Salmonoidea 179, 203
Salmonoidei 203
Salmostoma 144, 151
Sardina pilchardus 200
Sardinops sagax 232
Schismorhynchus 60, 62
Schultzidia 60
Sciaena hamrur 302

Scorpaenichthys marmoratus 231-232, 234
Securicula 151
Seriola lalandi 301
Sideria chlevastes 17, 22
Siluriformes 201
Solanaceae 49
Solanum 49

argentinum 49

elaeagnifolium 49

verbascifolium 49
Somniosus

microcephalus 89, 105

padficus 89, 105

(Somniosus) 105
Sparidarum rutoti 3 1 2
Sphecidae 27, 123

Sphex

figulus 125-126

fuliginosa 126

fuliginosus 123, 126-127
Sphyrna mokarran 89
Spirinchus 194
Spirobis 244
Squalus 228

acanthias 231-232
Stereolepis gigas 301
Styela 246-247

uniplicata 247
Styelopsis 246
Sundasalangidae 179, 181-182, 184-185, 189-190, 192,

194-195, 199, 201-202, 204, 211, 213, 218
Sundasalanx 179, 181, 185, 192-193, 198, 201, 204,
213-214,218

microps 170-181, 186, 191-192, 196, 198, 200-
202,209,213-214

praecox 179, 181, 186, 193, 196, 209, 213-214
Synodontidae 201

Tachysphex 27-42

acanthophorus 27, 29, 39-41

alayoi 29

apricus 27. 29-31, 34

arizonac 27, 29-30, 32, 35

armatus27, 29,40-41

ashmeadii 3 1

belfragei 36, 41

bohartorum 27, 29-31

brevicornis 29

brulli group 28-29, 36, 39-40

crenulatus 35-36

fulvitarsis 29

glabriorll, 36

idiotrichus 27, 29, 31, 34, 36

irregularis 27, 29, 31-32, 34, 36

julliani group 28-29

krombeini 34

krombeiniellus 27, 29, 41

lamellatus 27, 29-30, 32, 35-37

maurus 4 1

menkei 27, 29, 41-42

mirandus 27, 29, 32-34

mundus29, 36,40-41

musciventris 27, 29, 33-35

occidentalis 27, 29, 34-35

papago 27, 29, 35

pechumani 30-3 1

pompiliformis group 28-29, 31,36

psilocerus 30, 35

semirufus 33-34

Solaris 27, 29, 35

sonorensis 32, 37

spatulifer 27, 29, 35-36

spinulosus 40

INDEX

329

tarsatus 34

terminatus group 28-29

texanus 30

undescribed species 33

verticalis 27, 29, 31,36-37

yolo 27, 29, 37-39

yuma 27, 29, 39
Taeniura 284

grabata 285
Tetraodon 2,15

modestus 2
Tetraodon

(Arothrori) modestus 7-8

modestus 2

naritus 2

Tetraodontidae 1, 14-15
Tetraodontiformes 14
Thryssocypris 141-158

smaragdinus 141-158

tonlesapensis 141-158
Thunnus

albacares 301

obesus 301

Thysanopoda pect inata 108
Topobea 269-270, 281-282

brenesii 270

calophylla 269, 280-282

durandiana 282

elliptica 269-270, 281

pittieri28l
Trichoptera 14
Trygon 287, 295

margarita 287, 290

ukpam 295

Trypanorhynchida 110
Trypoxylon 125-126

apicale 126-127

apicalis 126

fieuzeti 127, 131

figulus 123-140

figulus barbarum 123, 127, 131

figulus koma 123, 132

figulus major 1 26

figulus forma major 126
figulus var. mo/or 1 36
figulus var. majus 1 26
figulus media 1 36
figulus forma media 136
figulus var. media 136
figulus medium 132, 136
figulus var. medium 1 36
figulus minor 1 32
figulus forma minor 132
figulus var. minor 132
figulus minus 132, 136
figulus var. minus 1 32
figulus minus var. rwZ?/ 1 36
figulus yezo 123, 127
WO/MS 123, 127
medium 123-140
mmus 123-140
rwfci 123, 139

Umbridae217

Urogymnus 283-284, 286-287, 295

africanus 285

asperrimus 285
Urolophoides 284
Uropterygius 22

alboguttatus 17, 22

kamar 22

marmoratus 17, 22-23

xanthopterus 17, 22

Verbesina encelioides 49

Xenopterus 1-3, 9

bellengeri 2

naritus 2-3
Xestotrachelus 43-58

hasemani 44, 56-57

, 48, 50, 55-57

Zalophus californianus 77-85, 229-230
Zoniopoda robusta 56

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